THE

NATURAL
HISTORY
MUSEUM

VOLUME 26 NUMBER 2 28 NOVEMBER 1996

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

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

Editor of Bulletin: Ms M.J. Short

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 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. All papers submitted for publication are subjected to external peer review for
acceptance.

A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn.
Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back
numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be
sent to:

Intercept Ltd.

P.O. Box 7 16

Andover

Hampshire SP 10 1YG

Telephone: (01264) 334748

Fax:(01264)334058

Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6
months for the UK, and 9 months for the rest of the world.

World List abbreviation: Bull. nat. Hist. Mus. Lond. (Bot.)
© The Natural History Museum, 1996

Botany Series
ISSN 0968-0446 Vol. 26, No. 2, pp. 75-2 1 7

The Natural History Museum

Cromwell Road

London SW7 5BD Issued 28 November 1996

Typeset by Ann Buchan (Typesetters), Middlesex

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

Bull. not. Hist. Mus. Land. (Bot.) 26(2): 75-217 Issued 28 November 1996

Studies in the genus Hypericum L. (Guttiferae)

6. Sections 20. Myriandra to 28. Elodes

THE NATURAL
HISTORY MUSEL

NORMAN K.B. ROBSON

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

.PRESENTED

CONTENTS

Introduction 76

Separation of sects Adenosepalum and Humifusoideum from the rest of Keller's sect. Euhypericum 76

Circumscription of seel. Adenosepalum 76

Sect. 20. Myriandra 76

Subdivision 76

Characters and variation 78

Leaves 78

Inflorescence 78

Flowers and fruits 79

Cytology and hybrids 79

Distribution and evolution 79

Sects 21. Webbia and 22. Arthrophyllum 82

Characters and variation 82

Distribution and evolution 83

Sects 23. Triadenioides, 24. Heterophylla and 25. Adenotrias 83

Characters and variation 83

Distribution and evolution 84

Sect. 26. Humifusoideum 85

Characters and variation 85

Distribution and evolution 87

Sects 27. Adenosepalum and 28. Elodes 88

Evaluation of sect. Elodes 88

Characters and variation 89

Morphology and subdivision 89

Cytology and hybrids 90

Distribution and evolution 90

Systematic treatment 92

Sect. 20. Myriandra (Spach) R. Keller 92

Sect. 21. Webbia (Spach) R. Keller 133

Sect. 22. Arthrophyllum Jaub. & Spach 137

Sect. 23. Triadenioides Jaub. & Spach 141

Sect. 24. Heterophylla N. Robson 146

Sect. 25. Adenotrias (Jaub. & Spach) R. Keller 147

Sect. 26. Humifusoideum R. Keller 153

Sect. 27 '. Adenosepalum Spach 170

Sect. 28. Elodes (Adans.) W. Koch 208

References 212

Systematic index 214

SYNOPSIS. Following citation of the type of a new subspecies (H. silenoides subsp. minus N. Robson) omitted from Part 8, those
sections of Hypericum directly related to sect. 1. Campylosporus that were not treated in Part 3 are considered, as well as sects
22. Arthrophyllum and 28. Elodes, which are closely related respectively to sects 2 1 . Webbia and 21 .Adenosepalum. A discussion
of the morphology, chromosome numbers, hybrids, distribution and evolution of each section is followed by a systematic account
of the 8 1 species in total that they contain.

Sect. 20. Myriandra is divided into five subsections: subsect. Centrosperma R. Keller, subsect. Pseudobrathydium R. Keller,
subsect. Suturosperma R. Keller, subsect. Brathydium (Spach) R. Keller and subsect. Ascyrum (L.) N. Robson stat. nov., all
additional to or with circumscriptions different from those inAdams's 1962 paper; and//, tenuifolium Pursh replaces//, reductum
W.P. Adams.

Sect. 27. Adenosepalum is divided into four subsections: subsect. Aethiopica N. Robson, subsect. Pubescentes N. Robsen,
subsect. Caprifolia N. Robson and subsect. Adenosepalum; and H. joerstadii Lid is treated as a hybrid, H. glandulosum x

reflexum.

© The Natural History Museum, 1996

76

INTRODUCTION

This part (Part 6) of the Hypericum monograph includes treatments
of the remaining sections directly related to the mainly African Sect.
1. Campvlosporus (sects 20-28), sects 2-3 having been treated in
Part 3 (Robson, 1985) and sects 29-30 in Parts 7 and 8 (Robson,
1987, 1990). Parts 4 and 5 will include the remaining sections, all
directly related to the Asian sect. 3. Ascyreia (sects 7-19).

Before turning to the main concern of this part, however, it is
necessary to remedy a nomenclatural omission in Part 8. The type of
Hvpericum silenoides subsp. minus, endemic to the Galapagos
Islands (Robson, 1990: 90), should have been cited as:

Ecuador, Galapagos Islands, Albermarle Island [Isabella], 825 &
945 m, 28 August 1905 (fl), Stewart 2064 (BM!-holotype; GH!, K!,
MO!-isotypes).

Separation of sects Adenosepalum and
Humifusoideum from the rest of Keller's sect.
Euhypericum

Sects 20-28 fall into six quite distinct groups, each related directly to
sect. 1 .Campvlosporus: (i) 20.Myriandra, with 2-5 styles ±appressed,
stamens apparently not fasciculate and black glands absent; (ii) 21 .
Webbia and its derivative 22.Arthrophyllum, with 3 styles spreading
from non-contiguous bases, stamens '3'-fasciculate, black glands
absent or present and leaves ± broad with densely reticulate venation;
(iii) 23. Triadenioides, with 3 styles spreading from contiguous
bases, stamens '3' -fasciculate, black glands present only in the most
derived species and leaves broad (but without clearly reticulate
venation) to± microphyllous; (iv) 24.Heterophylla and 25Adenotrias,
with3 styles spreading from contiguousbases,stamens'3'-fasciculate,
black glands absent and leaves microphyllous; (\)26.Humifusoideum,
with 3-5 styles spreading from contiguous bases, '3'-5 rather
indefinite stamen fascicles, black glands usually present and leaves
broad; and (vi) 21. Adenosepalum and its derivative 28. Elodes, with
3 styles spreading from contiguous bases, '3'-fascicled stamens,
black (or red in sect. 28) glands present and leaves broad. The
distributions of these groups are: Group (i) Eastern N. America,
eastern Mexico, Belize, Guatemala, Honduras Republic, Greater
Antilles, Bahamas, Bermuda; (ii) Macaronesia, eastern Mediterra-
nean; (iii) Socotra, S WTurkey, Levant; (iv) NWAfrica, Mediterranean,
NW Turkey; (v) New Guinea, Luzon, Taiwan, Sumatra, Java, S.
Africa to Ethiopia and Equatorial Guinea, Madagascar; (vi)
Macaronesia, Africa, W. Arabia, Europe, W. Caucasus, W. & S.
Turkey, Levant. Note that tropical and east Asiatic species hitherto
included in sect. Adenosepalum belong in fact to sect. Hypericum
sensu lato (see below).

Although each of these groups is easily distinguishable from the
others, members of sects 26 (Group v) and 27 (Group vi) have often
been included in a broad sect. Euhypericum Boiss., for example by
Keller (1925: 177). Keller placed such species in his subsect.
Homotaenium, by far the largest subsection in the section, which
contained most of the species with narrow continuous longitudinal
vittae on the capsule valves. Attempts to rationalize this heterogene-
ous agglomeration of species were made by Stefanoff (1932-34)
and Kimura (1951), both of whom divided it into sections, and by
Gorschkova (1949), who retained Keller's subsect. Homotaenium
but divided it into 15 series. In Part 1 of this work (Robson, 1977a),
I distributed Keller's species among sections 8, 9, 14, 17, 18, 26 and
27. Sects 17. Hirtella and 18. Taeniocarpium are distinguished
among these by having no regular intramarginal row of black glands
on the leaves and seeds with a smooth to papillose testa; in sect. 8.

N.K.B. ROBSON

Bupleuroides the leaf pairs are connate and glabrous and the styles
basally appressed; and in sects 9. Hypericum and 1 4. Oligostema the
petals become (roughly) erect after flowering, not tightly twisted
round the developing ovary as in sects 27. Adenosepalum and 28.
Elodes. Sect. Adenosepalum is therefore distinguishable among
species of 'sect. Homotaenium Boiss.' by the combination of free
leaf pairs with inframarginal black glands (or if leaf pairs are
connate they are pubescent), petals twisting round the developing
fruit, 3 distinct stamen fascicles (cf. sect. 26. Humifusoideum) and
seeds with a linear-reticulate to scalariform testa. The presence of an
indumentum in many species distinguishes these from members of
sects 9 and 14, whilst the widespread occurrence of flat-topped sepal
marginal glands in sect. Adenosepalum is paralleled only in some
species of sect. 17. Hirtella. Sect. 28. Elodes is differentiated from
sect. Adenosepalum principally by the floral modifications towards
specialized insect pollination, but also by the red marginal sepal
glands, the ribbed-scalariform seed testa and almost always by the
absence of inframarginal black leaf glands.

Circumscription of sect. Adenosepalum

When the foregoing criteria were considered, it became clear to me
that the Himalayan, south Indian and Chinese species that I origi-
nally included in sect. Adenosepalum (Robson, \911b) do not
belong there. The putative 'linking' species, the Himalayan H.
elodeoides Choisy, is in fact closely related to H. hengshanense W.T.
Wang, from SE China (Jiangsi, Hunan, Guangxi, Guangdong).
Indeed, Li (1990: 6) brought these two species together in his key to
Chinese Hypericum, placing both in sect. Adenosepalum.

Three glabrous, western Asiatic species allocated to sect.
Adenosepalum in Part 1 and previously in Flora of Turkey (Robson,
1 961 b) must also be excluded and transferred to sect. 1 2. Origanifolia.
These are H. huber-morathii N. Robson, H. minutum Poulter and H.
formosissimum Takht. They were originally excluded from sect.
Origanifolia because their capsules lacked the swollen vesicles
characteristic of H. origanifolium Willd. and H. aviculariifolium
Jaub. & Spach. H. minutum and H. formosissimum, however, have
interrupted vittae, which are also found in H. imbricatum Poulter
and H. salsugineum N. Robson & Hub.-Mor., both clearly related to
H. aviculariifolium; and H. minutum is clearly closely related to H.
huber-morathii. When other characters, such as glandularity, leaf
and flower colour and habitat (all in limestone rock crevices), are
taken into consideration, there can be little doubt that their correct
position is near H. aviculariifolium.

With the tropical and east Asiatic and Anatolian species excluded,
sect. Adenosepalum can be seen to have its primitive, shrubby
species in Macaronesia and three derivative groups: (i) African,
wholly glabrous; (ii) NW Africa and the Canary Islands to (a)
Somaliland and western Arabia and (b) the western Mediterranean,
wholly with indumentum; (iii) Europe to western Caucasus and
adjacent Turkey, the Mediterranean and Ethiopia (with adjacent NE
Sudan and Arabia) to northern Tanzania, nearly always with
indumentum. H. elodes L. (sect. 28. Elodes) is derived from group
(ii) and H. afrum Lam. (from Tunisia and adjacent Algeria) fits well
into group (i) rather than in sect. 9. Hypericum, where I originally
located it (Robson, 1977a).

Sect. 20. Myriandra

Subdivision

This well-delimited section comprises 29 species and 4 subspecies
that are mostly shrubs or wiry shrublets but include a few perennial

STUDIES IN HYPERICUM

77

Sect. 20 Myhandra

Relationships

7. tenuifolium
18

1 1 . brachyphyllum
18

21 adpressum

22. ellipticum

18

8. lloydii

19. microsepalum
18

9c. nitidum ssp. exile 13. fasciculatum

20. sphaerocarpum

9b. nitidum ssp.
nitidum

SUTUROSPERMA
16. apocynifolium 17. nudiflorum

29c. hypericoides
ssp. prostratum

29b. hypericoides

ssp. multicaule

9a. nitidum ssp. cubens
5. densiflorum

PSEUDO-
BRATHYDIUM
15. buckleyi

. tetrapetalum
18

27. edisonianum

24. dolabriforme
/ 18

BRATHYDIUM

23. myrtifolium
18

28. suffruticosum
18

4. lobocarpum
18

CENTROSPERMA

3. kalmianum
18

1 . frondosum

18,36

25. crux-andreae
18

29a. hypericoides
ssp. hypericoides
18

Fig. 1 Sect. 20. Myriandra. Relationships and chromosome numbers (2n) of the 29 species. Limits of the 5 (named) subsections indicated by bold lines.

herbs. They all lack black glands and have stamen fascicles closely
merged, so that the androecium appears to be polyandrous and (in
contrast to sect. 30. Trigynobrathys} remains so even in the smallest
flowers. The styles are slender and closely mutually appressed in the
more primitive species; but they have a tendency to part in fruit in
more advanced 4-petalled ones ('Ascyrum"). The stigmas are small.
Keller (1893, 1925) divided the group treated here as sect.
Myriandra into three main subgroups, viz. Ascyrum and two sec-
tions ofHypericum, with each section of Hypericum containing two
subsections:

Ascyrum - sepals and petals 4; placentation parietal; styles 2-3(4).
Hypericum - sepals and petals 5; placentation axile to parietal.

Sect. Mvriandra - stamens deciduous; placentation axile to parietal; styles

3-5.

Subsect. Centrosperma - placentation axile, pyramidal.

Subsect. Suturosperma - placentation parietal.
Sect. Brathydium - stamens persistent; placentation parietal or rarely

axile; styles 3.

Subsect. Eubrathydium - placentation parietal.

Subsect. Pseudobrathydium - placentation axile, pyramidal.

Following the inclusion of the American species of Ascyrum in sect.
Myriandra (Adams & Robson, 1961), Adams (1962) recognized
only one section, subdividing it into two subsections:

Subsect. Centrosperma - shrubs; leaves and sepals with articulation or
groove at base; sepals 5(4), deciduous; petals 5(4); stamens decidu-
ous. 15 species, i.e. Spp. 1-14 in the present treatment + 23. H.
myrtifolium.

Subsect. Pseudobrathydium - shrubs and perennial herbs; leaves and
sepals without articulation or groove at base; sepals 5-4, persistent
(deciduous in H. nudiflorum); petals 5^4; stamens persistent (except
in H. nudiflorum and H. apocynifolium). 14 species, i.e. Spp. 15-22,
24-29 in the present treatment.

From these attempts at infrasectional division it would appear that
there are three major groups (as Keller proposed), but that four
species (H. buckleyi, H. nudiflorum, H. apocynifolium and H.
myrtifolium) fall awkwardly across the proposed group divisions. In
addition, the Ascyrum group is heterogeneous, as Spach ( 1 8366) had
already realized. Most species have two unequal pairs of broad
appressed sepals, but in 19. H. microsepalum they are equal, narrow
and recurved. Spach placed the latter in a monotypic genus,
Isophyllum, as/, drummondii; and it is clearly not related to the other
4-petalled species but to 18. H. cistifolium, from which it appears to
have been derived.

The suggested interrelationships of species in sect. Myriandra are
shown in Fig. 1 , along with the five subsections into which they have
been divided here. It will be seen that there are two basal species (1 .

78

N.K.B. ROBSON

H. frondosum and 2. H. prolificum, both variable, especially the
latter), from which the remaining species have been derived. H.
frondosum is directly related to the Ascyrum group (Spp. 25-29) and
the Brathydiutn group (Spp. 23-24), whereas the Suturosperma
group (Spp. 1 6-22) and//, buckleyi (Sp. 1 5) are directly related to//.
prolificum (as are Spp. 3-14).

From Fig. 2, which shows the limits of certain characters, it will
be clear why Keller and Adams had difficulty in defining their
subdivisions and why no hard and fast lines can be drawn between
Myriandra sensu stricto (Spp. 1-14) and the remaining species. The
groups outlined above, however, can be treated as five reasonably
well defined subsections:

1 . Subsect. Centrosperma (p. 94) - Sepals and petals 5; sepals and
stamens deciduous; leaves and sepals articulated; sepals very
unequal to subequal; styles (2)3-5(6); placentation incompletely
axile to parietal; shrubs. Spp. 1-14.

2. Subsect. Pseudobrathydium (p. 1 1 2) - Sepals and petals 5; sepals
and stamens persistent; leaves and sepals not articulated; sepals
subequal; styles 3; placentation incompletely axile; low shrub.
Sp. 15.

3. Subsect. Suturosperma (p. 113) - Sepals and petals 5(4-3);
sepals and stamens deciduous or persistent; leaves and sepals not
articulated, sepals unequal or usually subequal; styles 3(4);
placentation incompletely axile to parietal; shrubs or perennial
herbs. Spp. 16-22.

4. Subsect. Brathydium (p. 122) - Sepals and petals 5; sepals
persistent, stamens deciduous or persistent; leaves articulated
incompletely or not at all, sepals not articulated; sepals very
unequal to subequal; styles 3(4); placentation incompletely axile
to parietal; shrubs or subshrubs. Spp. 23-24 (This subsection can
be distinguished from the preceding one by the unequal and
persistent sepals, more numerous stamens and widely branched
inflorescence, see key p. 93).

5. Subsect. Ascyrum (p. 124) - Sepals and petals 4: sepals and
stamens persistent; leaves articulated or not, sepals not articu-
lated; sepals very unequal; styles 2-3(4); placentation parietal;
shrubs or wiry shrublets. Spp. 25-29.

Characters and variation

(Fig. 2)

The species in sect. Myriandra vary from bushy shrubs up to 3 m tall
(1 . H. frondosum) to minute prostrate mat-forming wiry herbs (29c.
H. hypericoides subsp. prostratum). As mentioned above they lack
black glands, and the pellucid glands are always punctiform. The
contraction of the androecium to apparent polyandry is accompa-
nied by trends towards a reduction in number of members of the
floral whorls.

LEAVES. The most closely related species to sect. Myriandra in
sect. 1. Campylosporus is the north-east African H. synstylum,
which has pinnate leaf venation. Thus the venation in sect. Myriandra
is also basically pinnate. In the primitive, broad-leaved species (e.g.
1 . H. frondosum) the tertiary reticulation is very dense and clear, but
it is less clear or obscure in species with thicker leaves (e.g. 1 8. //.
cistifolium). The more advanced species in subsect. Centrosperma
have linear, often needle-like leaves with only a midrib visible
beneath.

The occurrence of a groove where the leaf joins the stem ('leaves
articulated') is a plesiomorphic character in Hypericum in general
and in sect. Myriandra in particular (Fig. 2). It is constant in subsect.
Centrosperma and present in primitive members of subsects

Suturosperma (Spp. 16, 17), Brathydium (23. H. myrtifolium) and
Ascyrum (Spp. 25, 26) but is wholly absent from subsect.
Pseudobrathydium ( 15. //. buckleyi). This character change is asso-
ciated with a tendency for the point of leaf-fall to move from the
basal groove to the petiole or pseudopetiole. Delayed basal leaf-fall
is found in 16. H. apocynifolium, 17. H. nudiflorum, 25. H. crux-
andreaeand 26. H. tetrapetalum, while suprabasal leaf-fall occurs in
15. H. buckleyi, 18. H. cistifolium, 19. //. microsepalum and the
remainder of subsect. Ascyrum (Spp. 27-29).

In the linear- to acicular-leaved species, there is a contrast be-
tween those in which the margin is revolute but the rest of the lamina
unaltered (12. H. lissophloeus and the H. galioides and //. nitidum
groups, Spp. 6-1 1) and the H. fasciculatum group (Spp. 13, 14), in
which the midrib area is raised beneath, forming a groove on each
side between it and the revolute margin (cf. Fig. 2).

INFLORESCENCE. The inflorescence of the near-ancestral //.
synstylum is 1-2-flowered, a state that is reflected in the 1-3-
flowered basic inflorescence of sect. Myriandra (usual in 1 . //.
frondosum). Two evolutionary tendencies lead to (i) the involve-
ment of an increasing number of nodes in flower-bearing
(basipetal) and (ii) an increasing degree of cymose (i.e. dichasial/
monochasial) branching (acropetal). Basipetal branching is domi-
nant in subsect. Centrosperma, leading to the long narrow
inflorescences of H. galioides, H. nitidum, H. lissophloeus and
their relatives. Acropetal branching is dominant in subsects
Suturosperma, Brathydium and Ascyrum, where involvement of
more than three nodes is rare, occurring only in forms of 25. H.
crux-andreae and 29. H. hypericoides. Pseudo-dichotomous
branching is confined to subsect. Ascyrum, where it occurs wholly
or partly in all species.

FLOWERS AND FRUITS. References have been made above to the
evolutionary reduction in number of floral members in sect.
Myriandra. The polyphyletic reduction from pentamery to tetramery
in the perianth is associated in subsect. Ascyrum with a reduction to
a dimerous gynoecium. The gynoecium is otherwise regularly
trimerous except in Spp. 2-5, where there appear to have been
secondary increases to 4-5-mery, and occasionally in the galioides,
nitidum and fasciculatum groups (Spp. 6-14).

The sepals are primitively unequal (e.g. in 1 . //. frondosum) and
remain markedly so in tetramerous forms of that species and in the
directly related subsect. Ascyrum, where the outer pair is large and
appressed (concealing the developing fruit) and the inner pair very
small or even absent (in some forms of 28. H. suffruticosum). In the
rest of the section except subsect. Brathydium, the inequality gradu-
ally becomes less, so that in the other species with a usually
tetramerous perianth, 19. H. microsepalum, the sepals are almost
equal. In subsect. Brathydium (Spp. 23, 24), however, both species
have unequal sepals, although the inequality is less in 24. H.
dolabriforme than in 23. H. myrtifolium. The curved-dolabriform
(hammer-shaped) petals in the former are the most extreme in form
in the section. Otherwise the petals do not provide specifically
distinct characters. The trend from deciduous to persistent stamens
has already been mentioned above, and the stamens otherwise vary
only in number and length, decreasing from c. 650 of up to 12 mm
long ( 1 . //. frondosum) to c. 30 of 2.5-4 mm (28. H. suffruticosum).
The relatively large number of stamens is one of the reasons for
associating//, dolab riforme with H. myrtifolium rather than with 20.
H. sphaerocarpum.

The ovary placentation is never truly axile, i.e. the placentae never
meet in the middle of the ovary. The primitive state, incompletely
axile or pseudo-axile, is found in Spp. 1-5 of subsect. Centrosperma
and in subsect. Pseudobrathydium (Sp. 15) and 23. H. myrtifolium.

79

Sect. 20 Myriandra

Characters

Inflorescence branching mainly basipetal \ Inflorescence branching mainly acropetal

Stamens deciduous

Fig. 2 Sect. 20. Myriandra. Limits of certain characters. Note the isolated apomorphic occurrences of 5(6)-styled and persistent stamens respectively.

Elsewhere there are parallel developments to truly parietal
placentation in all subsections. The variation in testa pattern of the
seeds is not so great as it is in some other sections: from reticulate to
finely scalariform via scalariform-reticulate and linear-foveolate.

Cytology and hybrids

(Fig. 1)

The chromosome number in sect. Myriandra is almost consistently
n = 9, 2n = 18, except for a record of 2n = 16 for 22. H. ellipticum,
a tetraploid record for 1 . H.frondosum (p. 96) and a population of 5.
H. densiflorum from Macon Co., North Carolina that had 2n = 27, n
= 1 1-14 (Adams, 1959). Despite this relative uniformity of chromo-
some number, natural hybrids appear to be very rare except in
cultivation, which suggests that the species are isolated by ecologi-
cal and/or biological factors rather than by geographical separation
alone. The cultivated hybrids, which arose spontaneously and doubt-
less continue to appear when suitable species are grown together, all
involve Spp. 1-6 only.

Distribution and evolution

(Fig. 3)

The north-east American and Caribbean sect. Myriandra is, as was
noted in Part 3 (Robson, 1985: 169), most closely related to those

species of sect. Campylosporus in which (i) the styles are com-
pletely united even in fruit and (ii) there are tendencies for the petals
to fall tardily (H. quartinianuni) and for the black glands to be
completely absent (H. synstylum). In these species, too, the develop-
ment of pinnate venation is almost complete, only the lowermost
pair of lateral veins or one of them remaining free. H. quartinianum
occurs from south-western Arabia (Yemen) to northern Malawi and
northern Mozambique, whilst the relatively apomorphic//. synstylum
is confined to two adjacent areas in eastern Ethiopia (Harar) and one
in northern Somalia.

There is a wide distributional gap, therefore, between north-east
Africa and south-eastern U.S.A., where 1. H.frondosum, the most
primitive species in sect. Myriandra, occurs sporadically from
Kentucky to Georgia and eastern Texas. There is a considerable
morphological gap, too, as H. frondosum has deciduous petals and
stamens, the stamen fascicles are completely united with a consider-
ably increased number of stamens, and the ovary is 3-merous. In the
leaves, the acute to obtuse apex of the African plants has become
apiculate-obtuse to rounded, and the venation is wholly closed and
pinnate, with widely spreading (not ascending) laterals, densely
reticulate tertiary venation and punctate (not elongate) glands.

From H. frondosum three clades diverge (i, vii, viii), these corre-

80

NC -»c Fla •» s Ala

II

Ga-> La

NC-»-n Fla-» e Tex

9c

w Cuba
nw Fla

9b

s SC, Ga,

n Fla, e Ala

NJ-+Ga, Ala-»Pa

e Tex,
e Okla -(• Ga?, SC

9a

w Cuba, Belize

N.K.B. ROBSON

Sect. 20. Myriandra

Distribution
14 nw Fla

13 se NC-*. Fla •» Miss

|9 s Ga, nw Fla

21

NC -*e La 18 Mass -*-Ga-

Tenn, Mo,

111, Ind

22

Minn-»Nfld
WVa, Tenn

16 Va, Tenn-»nw Fla 20

Ga' Fla Miss, [La], e Tex? lowa^Ohio^

La Ark n A^-a -*ne Tex, e Okla

17

c Fla

Mass -vSC-y
n Tex-*s Ohio

Que, Ont, Wis-*NY

26

s Ga, s Ala,
23 Fla, w Cuba

SC?, Ga-»Miss

29 b

s NC-»e La, c Fla

28

c Hisp (Dom R)

29a 29c

se NY-»Fla-»e Tex,

e Okla Va-*Fla-»-e Tex,

Okla-»Ky; e Mex-rHond;
Gr Ant, Jam; Bah, Berm; [Azores]

Fig. 3 Sect. 20. Myriandra. Distribution of the 29 species, showing major (==) and minor ( — ) disjunctions and trends (-»). Lower case letters indicate
compass points; square brackets indicate extinctions (Sp. 17) or introduction (Sp. 29c); roman figures indicate major clades (see text). Geographical
abbreviations used in Fig. 3: Ala - Alabama, Ark - Arkansas, Bah - Bahamas, Berm - Bermuda, Dom R - Dominican Republic, Fla - Florida, Ga -
Georgia, Gr Ant - Greater Antilles, Hond - Honduras Republic, 111 - Illinois, Ind - Indiana, Jam - Jamaica, Ky - Kentucky, La - Louisiana, Mass -
Massachusetts, Mex - Mexico, Minn - Minnesota, Miss - Mississippi, Mo - Missouri, NC - North Carolina, Nfld - Newfoundland, NJ - New Jersey,
NY - New York, Okla - Oklahoma, Ont - Ontario, Pa - Philadelphia, Que - Quebec, SC - South Carolina, Tenn - Tennessee, Tex - Texas, Va - Virginia,
Wis - Wisconsin, W Va - West Virginia.

spending to (i) H. prolificutn and the rest of subsect. Centrosperma
along with subsectsPseudobrathydium and Suturospertna, (vii) sub-
sect. Brathydium and (viii) subsect. Ascyrum. The transition from 1.
H.frondosum to 2. H. prolificum is gradual, consisting essentially of
reductions in size of parts and increase of variation north-westward
from NW Georgia to the uplands bordering the central Mississippi
basin; but there is a morphological 'gap' between these species, at
least in the wild. The wide distributional range of 2. H. prolificum
includes a comparably wide range of variation, so that as many as
five clades (ii-vi) appear to have arisen directly from it. 3. H.
kalmianum (ii) is a northern derivative in which reductions in size
(overall and of most parts) and inflorescence-branching are accom-
panied by a (secondary) increase to 5-mery in the ovary. The Great
Lakes type of distribution may be associated with the glacial nunatak
in this region, i.e. the species may have reached its present area in
pre-glacial times. Utech & Iltis (1970), however, favour the hypoth-

esis that H. kalmianum had a recent, post-glacial origin (from H.
prolificum). The other single-species derivative ( 1 5. //. buckleyi) (v)
has an area in the Carolinas and Georgia (the Blue Ridge Mountains)
that is wholly within that of//, prolificum. The speciating factors for
this dwarf straggling shrub with reductions in size and inflores-
cence-branching would seem to be altitude and exposure.

The remainder of subsect. Centrosperma (clades iii and iv) show
several parallelisms, so that extreme members of each (7. H.
tenuifolium and 8. H. lloydii on the one hand and 1 1 .//. brachyphyllum
on the other) have come to resemble one another, with resultant
taxonomic confusion. Adams ( 1 959, 1 962) resolved this confusion as
regards the U.S. species, and I have attempted to incorporate the taxa
from Cuba and Belize. Clade (iii) is relatively straight-forward. A
southward trend from //. prolificum resulted in two taxa, one north-
eastern (5. H. densiflorum), the other south-western (4. //.
lobocarpum). The reasons for treating these taxa as species rather

STUDIES IN HYPER/CUM

81

than subspecies are explained on p. 100. In both species there is a
reduction in size of flower and leaf and an increase in flower number.
In H. lobocarpum, these changes are accompanied by a marked
tendency towards a 5-merous ovary and lobed fruit, whereas in H.
densiflorum they are not. H. lobocarpum is mainly in the uplands to
the west of the lower Mississippi; H. densiflorum has a two-armed
distribution: coastal plain from New York to S. Carolina and along the
western side of theAppalachian Mts. But an 'arm' of//, lobocarpum^
area extends eastward in the south to south-eastern S. Carolina; and
where the southern end of the H. densiflorum area meets this 'arm',
in central Alabama, some intermediates (hybrids?) occur. A narrow-
leaved form of//, densiflorum found inTennessee (H. interior Small)
provides a morphological link to the narrower-leaved 6. H. galioides,
which has a coastal-plain distribution (N. Carolina to eastern Texas
excluding peninsular Florida). This species of relatively wet habitats
is ecologically distinct from its largely co-extensive reduced deriva-
tive, 8. H. tenuifolium, a plant of dry sandy habitats. Between the
'arms' of H. densiflorum'^ range there occurs 7. H. lloydii, a low,
spreading, narrow-leaved derivative of that species from the eastern
South Appalachian foothills. The remaining derivative clade of

subsect. Centrosperma (iv) forms the H. nitidum-H. fasciculatum
group, which comprises two subgroups: the H. fasciculatum group
(Spp. 12-14), confined to the American lowland mainland from N.
Carolina to Mississippi, and the H. nitidum group (Spp. 9-11),
having a similar distribution but extending into Louisiana, western
Cuba and Belize. The primitive forms of 9. H. nitidum would appear
to be in Cuba and Belize, but those of 13. H. fasciculatum are in
Florida. The overlapping areas and incomplete morphological differ-
entiation of the taxa in the H. nitidum group suggest that
land-connections between Belize and Florida via Cuba existed for a
long time.

The remaining clade from 2. H. prolificum (Clade vi) comprises
subsect. Suturosperma, of which the lowest branch (Spp. 16, 17)
includes species that are morphologically somewhat intermediate.
H. apocynifolium and H. nudiflorum, which have inflorescences that
are more acropetally branched than those of//, prolificum as well as
smaller flowers and broader leaves, form another east-west pair with
overlapping distributions: H. apocynifolium from Oklahoma and
Texas to Louisiana with outliers on the Florida-Georgia border, //.
nudiflorum from eastern Texas to Virginia excluding Louisiana,

Sects 21. Webbia and 22. Arthrophvllum

a) Relationships

b) Distribution and Characters

nanum var. prostratum

4b

vaccinii folium

5

pamphylicura 4d / ?

O nanum var. nanum

rupestre

s Turkey

canariense

revolutum ssp. revolutum

pal margin

ne trop. Africa

contiguous \ s«p«r»t«

style bases

Fig. 4 Sects 2 1 . Webbia and 22. Arthrophvllum. a) Relationships and chromosome number (2n). b) Distribution and limits of certain characters. For key to
annotations see Fig. 3 (p. 80).

82

N.K.B. ROBSON

Sects 23. Triadenioides, 24. Heterophylla. 25. Adenotrias

Relationships

aegypticum ssp. aegypticum
aciferum I C

aegypticuw
ssp. webbii 1 L

aegypticum ssp. maroccanum
20

dogonbadanicum N

I
ssp. smithii

socotranum

Fig. 5 Sects 23. Triadenioides, 24. Heterophylla and 25. Adenotrias. Relationships within sections and with species of sect. 1 Campylosporus, showing
chromosome numbers (2n).

where it is apparently extinct. The rest of the clade consists of a
southern branch (18. H. cistifolium in the Coastal Plain from N.
Carolina to Louisiana including Florida, 19. H. microsepalum in
Florida) and a northern one (20. H. sphaerocarpum in the central and
upper Mississippi valley 'giving rise to' the herbaceous 21. H.
adpressum (north-eastern) and 22. H. ellipticum (northern).

To return to the clades directly related to 1. H. frondosum, in
subsect. Bmthydium 23. H. myrtifolium (Clade vii) has smaller but
broad leaves and a more acropetally developed, more widely branch-
ing inflorescence than H. frondosum, and its distribution (Georgia to
Mississippi) is to the south-east and distinct. The derivative 24. H.
dolabriforme, with narrower leaves and more unequal sepals, has a
small relict area to the north-west centred in Kentucky andTennessee.
Finally, in Clade viii (subsect./UcyrM/n) the variable H. crux-andreae
occupies a wide area southward of a line from south-eastern Okla-
homa to New York (Long Island).Three of the derivative species occur
wholly (27. H. edisonianum, 28. H. suffruticosum) or largely (26. H.
tetrapetalum) within that area, H. tetrapetalum being found also in
western Cuba. The very variable 29. H. hypericoides, however, has a
much wider distribution and is divided into three subspecies. The
subspecies morphologically nearest to H. crux-andreae, 29a. subsp.
hypericoides, occupies almost the same region in the U.S.A. as that
species (Delaware and Maryland west to eastern Oklahoma and

eastern Texas), but it also occurs disjunctly along the Eastern Cordillera
in Mexico, Guatemala and the Honduras Republic. In addition, it is
found in the main islands of the Greater Antilles and in the Bahamas
and Bermuda. A record from the Azores suggest that it is a relatively
recent arrival there; but whether it came by natural extension of range
or with human assistance is not clear. Overlapping the northern
margin of the range of the erect 29a. subsp. hypericoides is that of the
more spreading 29b. subsp. multicaule (Oklahoma to Massachusetts);
and in Hispaniola (Dominican Republic) there is a prostrate, almost
herbaceous form (29c. subsp. prostration).

Sects 21. Webbia and 22. Arthrophyllum

Characters and variation

(Fig. 4a, b)

Sects Webbia and Arthrophyllum form a monophyletic group di-
rectly related to sect. Campylosporus. As will be explained on p. 1 35,
I now regard the nearest relative of H. canariense (sect. Webbia) to
be not//, roeperianum (as in Robson, 1985: 166-168, ff. 1-3) but//.
revolutum subsp. revolutum, more specifically the relatively broad-
leaved form of that species that occurs sporadically in Ethiopia.
From it, H. canariense differs inter alia in having '3' stamen
fascicles, a 3-merous ovary with divergent styles, no black glands,

STUDIES IN HYPERICUM

Sects 23. Triadenioides. 24. Heterophvlla. 25. Adenotrias

Characters

83

pre

Black or r

discolor

basally narro
or petiola

Fig. 6 Sects 23. Triadenioides, 24. Heterophvlla and 25. Adenotrias. Limits of certain characters.

and leaves with densely reticulate tertiary venation. In addition, the
bark is smooth and usually whitish grey, a character that links it to
the species of sect. 22. Indeed, apart from a difference in habit (trees
or ± erect shrubs up to 4 m versus low, compact and rounded to
prostrate shrubs up to c. 0.9 m), the only essential differences
between these sections would appear to be in the style bases (con-
tiguous then divergent v. separate) and testa sculpturing
(linear-foveolate v. minutely rugulose).

Within sect. Arthrophyllum there is a return to the presence of
hypericin-containing glands in 1 .//. rupestre and 2. H. pamphylicum,
black in 1 and red in 2, although in all other respects//, pamphylicum
is more advanced than H. rupestre, e.g. in its perfoliate leaves. In the
other branch of Fig. 4 there is, as well as a reduction in overall size
and in size of parts, a change from cordate to cuneate leaf-base.

Distribution and evolution

(Fig. 4b)

Sects Webbia and Arthrophyllum, though morphologically similar,
are geographically divergent H. canariense is confined to the
Canary Islands (except the drier eastern ones) and Madeira, thus
being separated from its apparent nearest ancestral relatives in
Ethiopia by the width of Africa. It is separated by the length of the
Mediterranean from sect. Arthrophyllum, which has an interrupted
distribution from Vilayet Antalya in south-west Turkey to the moun-
tains on the Lebanon-Syria border. The facts that (i) the five species

in this section are so morphologically and geographically disparate
and that (ii) each occupies a restricted area suggest that it is a
relatively ancient group that evolved (or at least diverged) in Tertiary
times; and the link with Macaronesia further associates it with the
pan-Mediterranean flora of that epoch.

Sects 23. Triadenioides, 24. Heterophylla and 25.
Adenotrias

Characters and variation

(Figs 5, 6)

Sects 23-25 all derive from the H. socotranum group of sect.
Campylosporus, sect. 23. Triadenioides having a Socotra-E. Medi-
terranean distribution and sects 24. Heterophylla and 25. Adenotrias
(as a group) an Atlantic-Mediterranean one.

Sect. Triadenioides is most closely related to H. socotranum
subsp. socotranum, and its species either share with that taxon a
narrowing of the leaf-base (2. H. scopulorum in part) or have a
petiole. The leaf venation is simply pinnate, a state which can be
derived from that of subsp. socotranum by elimination of the 1-2
pairs of basal veins and expansion of the (closed) midrib branching
to occupy the whole lamina. The flowers in sect. 23 are much
reduced in size from those of subsp. socotranum, and trimery
(ovary) and pseudotrimery (stamens) have developed. The three
more primitive (Socotran) species have discolorous leaves; but in the

84

N.K.B. ROBSON

Sects 23. Triadenioides. 24. Heterophylla, 25. Adenotrias

Distribution

s Turkey
(Antalya, Isparta)

sw Crete

Lebanon, w Syria,
s Turkey

s Turkey (Hatay)
w Syria

n Socotra

ne Libya (Derna)
1C

Ib

Lampedusa, Malta,
/ Sardinia,
Ionian Is. ,
sw Greece,
nw Crete

la

s Morocco, c Algeria
(s Atlas)

Socotra '

Fig. 7 Sects 23. Triadenioides, 24. Heterophylla and 25. Adenotrias. Distribution of the species and of the immediately related species in sect. 1
Campylosporus. H. tortuosum (Sect. 23, Sp. 3) also occurs in western Socotra (Map 21).

Mediterranean 4. H. ternatumand 5. H. pallensboth sides of the leaf
are glaucous, and in the former the leaves are 3-whorled. There is a
multiplication of flowers in the inflorescence in both groups, but red
or black glands are found only in the Mediterranean group.

The essential distinction between sects 24. Heterophylla and 25.
Adenotrias is the presence in the latter of specialized pollination and
dispersal adaptations (see opposite and p. 150; also Robson 1972,
1981;Reynaud, 1985).

Distribution and evolution

(Fig. 7)

Although there is a considerable morphological gap (especially
florally) between the large-flowered Socotran species of sect. 1 (H.
balfourii and H. socotranum) and all three derivative sections, the
respective geographical disjunctions are very different. Whereas
three disjunct but closely related species of sect. 23. Triadenioides
occur in one mountain range in Socotra (1. H. fieriense, 2. H.
scopulorum, 3. H. tortuosum, 1 and 2 being both endemic to it and
directly related to H. socotranum), there is then a morphological and
geographical gap in this section between them and the more highly
evolved 4. H. ternatum and 5. H. pollens in the north-east Mediter-

ranean. On the other hand, the main morphological and evolutionary
gaps in the other evolutionary line are between northern Socotra (H.
balfourii) and respectively north-west Anatolia (H. heterophyllum,
sect. 24. Heterophylla) and southern Morocco (the largest form of
the heterostyled H. aegypticum, la. subsp. maroccanum, sect. 25.
Adenotrias). H. heterophyllum is more advanced than H. aegypticum
in several characters (e.g. (i) elongate upper stem-internodes and
condensed lower stem-internodes, resulting in gemma-like axillary
buds; (ii) a several-flowered inflorescence; (iii) a chromosome
number of n = 9, not 10); but it lacks the floral features by which the
species in sect. Adenotrias are adapted for specialized insect pollina-
tion (heterostyly, etc.) and the carunculate seeds that aid them in
dispersal. H. heterophyllum is therefore most likely to have been
derived from a precursor of sect. Adenotrias, one in which the
specializations had not developed. ' The divergence of sects 24 and
25 can thus be seen as an east-west split in the Mediterranean region.
Within sect. 25 there is a very disjunct reduction trend in 1. H.
aegypticum from south Morocco along the southern slopes of the

' I am grateful to Chris Humphries for doing a cladistic analysis of sects Heterophylla
and Adenotrias that supported this conclusion.

STUDIES IN HYPERICUM

85

Sect. 26. Humifusoideum

Relationships

beccarii ssp. steenisii

(ii) nokoense

raacgregorii

3

saruwagedicuin 2
24

peplidifoliun

10 16

wilmsii

bif urcatuzn

Fig. 8 Sect. 26. Humifusoideum. Relationships and chromosome numbers (2n) of the 10 species. 6(i) H. nagasawai has 2n=36 (not shown).

Atlas Mts in Algeria to Lampedusa, the Maltese Islands and one
mountain in Sardinia. There is then a 'jump' to the Ionian Islands, a
small area in the western Peloponnisos and another in north-west
Crete, then another disjunction to the Derna area of Libya. The tall
erect long-leaved Moroccan plant gradually becomes small, decum-
bent to prostrate (in Crete) and short-lived, so that the two ends of the
morphocline look very different. It is, however, almost continuous,
so that no populations merit more than subspecific rank, and even
that rank may sometimes prove difficult to justify. There is a bigger
morphological and geographical disjunction from western Greece to
the northern Levantine 2. H. russeggeri, a species with leaves
broader above (not at or below) the middle and pedunculate several-
flowered inflorescences that was once also found in north-west
Turkey. The morphocline from this species then turns westward to
south-western Crete, where 3. H. aciferum resembles a smaller
prostrate-branched version of H. russeggeri.

With the possible exception of the Sardinian localities of H.
aegypticum subsp. webbii, all regions of the Mediterranean occu-
pied by members of sect. Adenotrias are strongly influenced by
maritime conditions; the plants typically grow among maritime
limestone rocks. How then are we to explain the distribution of H.
aegypticum subsp. maroccanum along the southern edge of the Atlas
ranges, facing the Sahara desert? There is apparently no evidence of

an ancient ocean in that area. Indeed, the traditional course of the
Tethys Sea (and hence of the boundary between Gondwanaland and
the northern supercontinent, Laurasia) is through the Strait of Gi-
braltar. If this boundary were to have taken a more southerly course,
so that theAtlas Massif 'belonged' to Laurasia, not to Gondwanaland
(as was proposed by e.g. Melville, 1967: 294), then the south face of
the Atlas Massif would indeed be part of the northern shore of the
Tethys Sea, and//, aegypticum would at one time have been growing
in typical maritime conditions.

As regards the Sardinian localities of H. aegypticum, these com-
prise several isolated sites where the species forms part of 'an
archaic relict flora no longer in balance with the present Mediterra-
nean macroclimate' and has presumably originated 'from the lands
of theTertiary Tyrrhenian-Iberian-Riffian continent' (Arrigoni, 1 965).

Sect. 26. Humifusoideum

Characters and variation

(Figs 8, 9)

Despite the long distance between the locality of the most primitive
species in sect. Humifusoideum (NE New Guinea) and those of its
nearest relatives in sect. Campylosporus (H. lanceolatum in Reunion
and H. madagascariense in Madagascar), the morphological gap is

86

N.K.B. ROBSON

Sect. 26. Humifusoideum

Characters

X pseudo-dichotomous

Inflorescence branching
dichasial

. 10 .

laterals
dense ,
reticulation
sparse
or absent

more than 3

laterals spaced,
reticulation *
dense

Inflorescence
branchina

Leaves

Leaf venation

Fig. 9 Sect. 26. Humifusoideum. Limits of certain characters. Note (i) the isolated apomorphic occurrences of more than 3 styles and the absence of black
glands and (ii) the development of pseudo-dichotomous and then (in Sp. 6) dichasial inflorescence-branching. Spp. 6(i) in part and 6(ii) also have
dichasial inflorescence branching, and both have black glands (not shown).

not so very great. H. sewense has an overall resemblance to the latter,
but it is smaller in stature and has three spreading styles (not five
appressed ones). Its venation of closely parallel lateral veins some-
times flanked by rows of streaks or dots is reminiscent of that of H.
lanceolatum subsp. angustifolium, and the similar more numerous
veins in the broader leaves of H. bifurcation are even more reminis-
cent of leaf- venation in that taxon. In the species with broader leaves
(Spp. 5-7a, 8-10) the laterals are more widely spaced with evident
tertiary reticulation, and in the secondarily narrow leaves of 7b. H.
beccarii subsp. steenisii and some forms of 6(i). H. nagasawai there
are remnants of this reticulation.

The occurrence of black glands is variable in sect.Humifusoideum,
but they are always present somewhere in the plant (on the anthers
and/or the margins of leaves, sepals and/or petals) except in (i) 3. H.
macgregorii and (ii) 6. H. pulogense and a few populations of its

nearest ancestral relative, 5. H. papuanum. Absence of black glands
is thus apomorphic in this section. When present they are never
laminar, except sometimes in the petals of 6(i). H. nagasawai, 6(ii).
H. nokoense and 7a. H. beccarii subsp. beccarii. Another reversal in
normal trend direction is in style number. Spp. 1^4- all regularly have
a 3-merous ovary, but in 5. H. papuanum the ovary varies from 3-
mery to 5-mery and even exceptionally 6-mery (Smith, 1 941 ); and in
6. H. pulogense it is occasionally 4-merous. There is a parallel trend
among the African species: 3-4(-5)-merous in 8. H. natalense, 3—4-
merous in 9. H. wilmsii and (4-)5-merous in 10. H. peplidifolium.
The fascicles of stamens are more or less obscure throughout the
section, but the increased number of styles is apparently accompa-
nied by a 'return' from '3' (i.e. 2+2+1) fascicles to 5 vaguely distinct
ones (Saunders, 1937). Keller's (1893, 1925) classification of H.
peplidifolium in a separate monotypic section because of its alleged

STUDIES IN HYPERICUM

Sect. 26. Humifusoideum

87

Distribution

(ii) Taiwan

111 Taiwan

Philippines (n Luzon)

6

nw Sumatra

7b

7a

w Sumatra, w Java

e Zimbabwe to Ethiopia;
s Sudan Rep.; Angola;
Cameroon, F. Poo

10

we to e New Guinea

e Zimbabwe, e Transvaal,
O.F.S. . Lesotho, e Cape Prov.
/ Central Madagascar

w central to
e New Guinea

8

Swaziland, a Natal,

ne Cape Prov. (Transkei)

ne New Guinea
( Madang)

ne New Guinea
(E. Highlands, Morobe)

Madagascar £ 1
Reunion

Fig. 10 Sect. 26. Humifusoideum. Distribution of the 12 species and the immediately related species in sect. 1 Campylosporus. For key to annotations see
Fig. 3 (p. 80). The gap in distribution between Sp. 6 (Luzon) and Spp. 6(i) and 6(ii) (Taiwan) constitutes a further small disjunction.

possession of 3' stamen fascicles and 5 styles does not appear to be
warranted.

Inflorescence branching is also variable in sect. Humifusoideum.
In 1 . H. sewense, as in the primitive species of sect. Campylosporus,
the flowers are nearly always solitary, terminating lateral branches.
In the only collection known so far, however, there are one or two
examples of pseudo-dichotomous branching. In the derivative nar-
row-leaved species (Spp. 2-3), branching is wholly lateral, but in 4.
H. bifurcation it is wholly pseudo-dichotomous. This type or the
derived 'pseudo-axillary' branching (see Sp. 10) is constant in the
rest of the section except in 5. H. papuanum, 6. H. pulogense, 6(i).
H. nagasawai and 6(ii). H. nokoense, where the solitary flower is
sometimes replaced by a regular dichasium or (Sp. 6) mixed dichasial/
pseudo-dichotomous branching.

Distribution and evolution

(Fig. 10)

The extraordinary distribution and a consequent theory of evolution
of sect. Humifusoideum were discussed in Part 2 (Robson, 1981:
212), so only a modified summary will be given here. The morpho-
logical and geographical relations between 1. H. sewense and the

Mascarene species of sect. Campylosporus imply that the ancestors
of sect. Humifusoideum spread to Australia (probably via Antarc-
tica) before the final separation of the African and Australian
land-blocks, about 140 m.y. B.P. When the Australian plate (which
includes New Guinea) made contact with the South-east Asian plate
(c. 40 m.y. B.P.,fide Audley-Charles, 1987), a land-connection was
made that could have allowed the H. papuanum group to migrate
from New Guinea into (i) Java, Sumatra and (ii) the Philippines and
Taiwan via the New Guinea track (Steenis, 1964; Smith, 1986).2 On
the other hand, the close relationship between the New Guinean 4.
H. bifurcatum and 5. H. papuanum and the south-east African 8. H.
natalense suggests that the early evolution of sect. Humifusoideun
took place in Australia at a time when contact with the African plate
was still possible. The occurrence of 9. H. wilmsii on both sides of
the Mozambique Channel (implying a long period of evolution of
sect. Humifusoideum in Africa) constitutes further evidence in fa-
vour of that hypothesis. Following this 're-entry' into Africa, there
would thus appear to have been a subsequent northward and west-

: The similarity between this migration 'track' and the distributions respectively of H.
papuanum, H. pulogense and H. beccarii (especially subsp. steenisii) is remarkable.

88

N.K.B. ROBSON

Sects 27. Adenosepalum, 28. Elodes

22 cuisinii

Relationship?

18c

annulatura
tap. afromontanum

annulatum
asp. intermedium

PUBESCENTES

caprifolium
18,16

CAPRIFOLIA
14

naudinianura
18

aathiopicum
map. aathiopicum

6b

16 [20, 32?]-? r«fl«xun
18

x joerstadii

glandulosum
18.40

Fig. 1 1 Sects 27. Adenosepalum and 28. Elodes. Relationships and chromosome numbers (2n) of the 25 species. Limits of (named) subsections of sect.
Adenosepalum and of sect. Elodes indicated by thin lines.

ward migration (Spp. 8-10) that also extended eastward to Mada-
gascar (Map 29 (p. 166) and Robson, 1958: 441, map 2).

It should be reiterated that, with the exception of some probably
wading-bird-dispersed species in sect. 30. Trigynobmthys (Robson,
1990: 1 1), there is practically no evidence for long-distance disper-
sal in Hypericum, so that the wide gaps in distribution in sect.
Humifiisoideum are very unlikely to have been achieved by such
means.3 On the other hand, vertical variation in land level in Aus-
tralia and New Guinea and increasing aridity in Australia both imply
that the New Guinea species reached their present montane refugia
relatively late (Audley-Charles, 1987; Morley & Flenley, 1987;
Robson. 1993fl). It is not possible, from present evidence, to decide
whether this happened before or after they died out in an increas-
ingly arid Australia.

' Pace Smith ( 1 977: 99), who placed a New Guinea montane Hypericum in a group with
'dust seeds weighing less than 0.1 mg which may also be predominantly wind-
dispersed' . Also see Robson ( 1993«), where the distributional history of all the tropical
montane groups of Hypericum is discussed.

Sects 27. Adenosepalum and 28. Elodes

Evaluation of sect. Elodes

From Fig. 1 1 (p. 88) it will be seen that sects 27 and 28 form a
monophyletic group, but that sect. 27. Adenosepalum alone is
paraphyletic. The justification for recognizing sect. Elodes is similar
to that which was adduced for separating sect. 25. Adenotrias from
sect. 24. Heterophylla: the development of a specialized pollination
syndrome (Robson 1972, 1981). In general the two syndromes are
very similar: the development of (i) stiffly erect sepals that confine
the petals, thus forming a pseudo-tubular corolla with a subrotate
'limb'; (ii) a non-secreting ligule on each petal, which guides the
tongue of a nectar-seeking insect; (iii) small bifid non-secreting
structures at the base of the ovary, which alternate with the 'three'
stamen fascicles and. like lodicules of grasses, swell to expand the
corolla; and (iv) union of all the filaments in each fascicle for about
0.7 of their length. H. elodes, however, lacks the heterostyly present
in sect. Adenotrias and the caruncles that adorn the seeds in that
section.

STUDIES IN HYPERICUM

89

When these specializations are discounted, however, H. elodes
can be seen as a development of the H. caprifolium group of sect.
Adenosepalum (Spp. 13-15). It shares with them a spreading
indumentum and humid habitats with an Atlantic climate, taking
these and other tendencies to extremes. Thus the indumentum
extends to the sepals, not only to the base of the inflorescence; the
testa sculpturing is ribbed-scalariform, not merely finely scalariform;
the inflorescence usually develops from one axil only of the pair; and
the habitat is aquatic, not merely moist or wet. In addition, H. elodes
shows evidence of a diminution of hypericin content in that the
inframarginal leaf glands are pale, the sepal marginal glands red and
the anther gland amber, whereas all these glands in 1 5. H. caprifolium
are black, as are all but the anther gland in 13. H. coadunatum and
1 4. H. naudinianum. In addition, the inframarginal black leaf-glands
in H. coadunatum are irregularly spaced or even absent. In leaf
shape, H. elodes is most similar to H. coadunatum, and its extreme
Atlantic habitat is more like that of the Canary Islands than of the
sub-Mediterranean areas of the other two species. The chromosome
number of n = 10 (Delay, 1972) (if correct) would, however, indicate
a separation of//, elodes from all three of the other species (n = 9, 8);
but my own (unconfirmed) count of 2n = 32 would provide evidence
for a derivation from within the H. caprifolium group. Support for

the latter theory was provided by Al-Bermani et al. (1993), who
recorded 2n = 16.

Characters and variation

(Figs 11, 12)

MORPHOLOGY AND SUBDIVISION. The species in sects
Adenosepalum and Elodes vary from bushy or straggling shrubs to
wiry or soft perennial herbs. They can be divided into three groups,
one wholly glabrous (Spp. 1-6) and two with indumentum (Spp. 7-
15 and Spp. 16-24). The two Macaronesian species (1. //.
glandulosum, 16. H. reflexum) are morphologically so distinct that
intermediates, e.g. H. glandulosum with some indumentum, are
assumed to be of hybrid origin (H. x joerstadii). Excluding these,
presence of indumentum provides a clear-cut distinction between
groups, except for forms of 1 7. H. montanum and 1 8a. H. annulatum
subsp. intermedium that appear to be secondarily glabrous. The two
indumentum-bearing groups can almost be differentiated by the
absence (Spp. 7-15) or presence (Spp. 16-24) of bracts with glandu-
lar auricles or densely crowded basal glands. Only in the shrubby 1 6.
H. reflexum and in 22. H. cuisinii and reduced forms of 23. H.
lanuginosum are these completely absent; but they occur independ-
ently in the bracteoles of 15. H. caprifolium, in which however the

Sects 27. Adenosepalum, 28. Elodes

Characters

Indumentum

linear-reticulate
to foveolate

scalariform
- reticulate

scalariform to • •
ribbed-scalariform

and-fringed

Fig. 12 Sects 27. Adenosepalum and 28. Elodes. Limits of certain characters. Note the isolated apomorphous occurrences of lack of indumentum.

90

N.K.B. ROBSON

pairs of leaves are united. If, in addition, the completely glabrous
inflorescence and free leaves of Spp. 16-24 are contrasted with the
pubescent inflorescence of Spp. 7-12 and the united leaf-pairs of
Spp. 13-15, a subdivision of sect. Adenosepalum into four subsec-
tions becomes possible:

1 . Subsect. Aethiopica (p. 172) - Plant completely glabrous; leaves
free; bracts and bracteoles not glandular-auriculate. Spp. 1-6.

2. Subsect. Pubescentes (p. 181)- Plant with indumentum up to the
sepals (Spp. 7, 9) or the lower part of the inflorescence (Spp. 1 0-
1 2) or rarely only to the base of the inflorescence (Sp. 8); leaves
free; bracts and bracteoles not glandular-auriculate. Spp. 7-12.

3. Subsect. Caprifolia (p. 189) - Plant with indumentum up to the
base of the inflorescence; leaves all or mostly connate in pairs;
bracts and bracteoles glandular-auriculate or not. Spp. 13-15.

4. Subsect. Adenosepalum (p. 193) - Plant with indumentum up to
the base of the inflorescence or rarely stems (Sp. 23 in part) or
leaves (Sp. 16) or wholly (Spp. 17, 18b, both in part) glabrous;
leaves free; bracts and bracteoles usually glandular-auriculate.

The occurrence of protruding marginal leaf-glands, which gave
rise to the epithet glandulosum for Sp. 1, appears to be a local
apomorphism. Although H. glandulosum is basic to the whole of
sect. Adenosepalum, these prominent leaf-glands have been ob-
served elsewhere only on one, particularly large-leaved specimen
of 7. H. pubescens from Morocco (Lewalle 9926, from Moyen
Atlas), apart of course from some specimens of Ix. H. x
joerstadii.

CYTOLOGY AND HYBRIDS. The ancestral group in sect. 1.
Campylosporus from which sect. Adenosepalum arose almost cer-
tainly had a basic chromosome number of x = 12, 11 or 10, most
likely 10 (see Robson, 1985: f. 1), but this number has been found
only in 1 . H. glandulosum (as 4x) and possibly in H. elodes (as 2x).
All the other counts from sects Adenosepalum and Elodes (except
for the problematic 2n = 20) are 2n = 18 or 36 (the latter in 7. H.
pubescens only) or 2n = 16 or 32. The trend has thus been x = 10, 9,
8 with occasional tetraploidy on each basic number (Fig. 11). No
count is recorded from subsect. Aethiopica other than 2n = 18, 40
from H. glandulosum; subsect. Pubescentes has 2n = 18, 36, 16;
subsect. Caprifolia has 2n = 18, 16; and subsect. Adenosepalum 2n
= 18, 16, 32. In sect. Elodes the recorded numbers are 2n = 16, 20,
32.

Natural hybrids apparently occur between 1 . H. glandulosum and

16. //. reflexum (H. x joerstadii) and between 7. H. pubescens and 9.
H. tomentosum (H. tomentosum var. intermedium). In cultivation,

17. H. montanum was successfully hybridized with H. tetrapterum
Fr. (= H. acutum Moench) and H. maculatum Cr., species in sect. 9.
Hypericum with the same chromosome number (2n = 16). The
fertility, however, was low 'and the plants (H. tetrapterum) and
seedlings (H. maculatum) were variegated (cf. Noack, 1 934; Robson,
1981: 171). Noack also produced hybrids between H. montanum
and H. perforatum L. (sect. Hypericum, 2n = 32), this time less
sterile but pentaploid, because of the apomictic ovules in H.
perforatum (Noack, 1939).

Distribution and evolution

(Fig. 13)

The most similar species to 1. H. glandulosum (Canary Islands,
Madeira) in sect. 1. Campylosporus would appear to be H.
quartinianum (SW Arabia, E. Africa) rather than the closely related
H. synstylum (Ethiopia, N. Somalia) suggested in Robson (1985)
(see p. 1 75). This distributional gap is comparable with that between
H. revolutum subsp. revolutum (sect. Campylosporus) and H.

canariense(stc\.. 2 1 . Webbia) (see p. 1 35). But, whereas H. canariense
is related to the eastern Mediterranean sect. 22. Arthrophyllum, the
relationships of the Macaronesian H. glandulosum and H. reflexum
are with the African mainland.

One, clearly ancient trend (i) gave rise to subsect. Aethiopica in
tropical and south Africa, with the relict Angolan 2. H. abilianum in
the west and all the other species in the east. These form two pairs,
a southern pair of subspecies (6. H. aethiopicum) with a disjunct
occurrence in Angola of the more highly evolved form of 6a. subsp.
sonderi, and a more northern pair of species (3. H. conjungens, 4. H.
kiboense) with an outlying derivative in NW Africa (5. H. afrum).

The second trend directly related to H. glandulosum (ii) com-
prises subsects Pubescentes and Caprifolia. In Pubescentes, 1. H.
pubescens is distributed from southern Morocco north to the south-
ern Iberian Peninsula thence eastward to western Libya, Malta,
Sicily, southern Italy? and Sardinia (extinct?). In the south of
Morocco, where it becomes reduced in habit and in size of parts, it
approaches the apparently derivative 8. H. psilophytum, with short
appressed stem-hairs. This species is confined to the Central Saha-
ran mountains of southern Algeria (Ahaggar, Tefedest and Tassili
n'Ajjer) and two localities in the southern Moroccan Atlas. The very
closely related 9.H. tomentosum occupies a generally more northern
area: northern Morocco and NW Algeria, Spain and central Portu-
gal, mediterranean France and adjacent Italy, and the larger islands
(Corsica - extinct?, Sardinia, Majorca). It is also recorded from
further east in Algeria and in Tunisia. In some parts where its
distribution overlaps that of H. pubescens, intermediates (hybrids)
occur (e.g. in SE Spain (Valencia), the Morocco-Algeria border area
and possibly Sardinia).

The other part of subsect. Pubescentes is separated from 7. H.
pubescens by the width of Africa. 10. H. somaliense (N. Somalia),
11. H. collenettiae (Saudi Arabia: Asir) and 12. H. sinaicum (Mt
Sinai and mountains east of the Gulf of Aqaba in Saudi Arabia) form
a geographical and morphological reduction series northward, each
population being small and isolated. According to Mrs Sheila
Collenette, her eponymous species may already be extinct.

Subsect. Caprifolia and sect. Elodes form a north-western group.
14. H. naudinianum (NW Morocco, adjacent Algeria) is morpho-
logically nearest to H. pubescens in most respects and 13. H.
coadunatum (Gran Canaria) is in others; so it seems likely that an
ancestral group diverged from H. pubescens (or its precursor) before
it in turn differentiated into a mainland and an island species. Each
then differentiated to form another species, the eastern Spanish 15.
H. caprifolium from H. naudinianum and H. elodes from H.
coadunatum; or perhaps H. elodes arose before Spp. 13 and 14
differentiated.

Subsect. Adenosepalum also has its primitive species in the
Canary Islands (16. H. reflexum), and its derivatives likewise split
into (north-) eastern and (south-) western groups. The western group
consists of only 17. H. montanum, which extends in distribution
from Morocco northward to northern England and central Scandina-
via and eastward to Poland, the Ukraine, Georgia and adjacent
Turkey but is absent from most of the Mediterranean peninsular
regions and the islands. This species can be separated from the
eastern 1 8. H. annulatum only by the denser inflorescence and/or the
distribution of indumentum. H. annulatum itself has a widely dis-
junct distribution. The form nearest to//, montanum occurs in Saudi
Arabia (Asir) and the adjacent African mainland, whence there is a
morphological trend through Ethiopia, Kenya and Uganda to north-
ern Tanzania. This can be split into a northern and a southern
subspecies ( 1 8a. subsp. intermedium and 1 8c. subsp. afromontanum
respectively) with a somewhat intermediate population in eastern
Ethiopia (Harar). A dwarf species related to the northern subsp.

STUDIES IN HYPERICUM

Sects 27. Adenosepalum, 28. Elodes

91

Distribution

r\ f\ Aegean Is. ,

•^ ^- w Turkey (Izmir)

s Turkey, Levant, Cyprus 23

Libya (Derna) 24

Aegean region, 2 |
sw Turkey

se Greece

(Euwoia, Andhros)

19

20

ne Greece,
Thasos,
Sanothraki

Egypt (Sinai) ,
nw Saudi Arabia

I8a.

sw Saudi Arabia, ne Sudan,
Eritraea, n Ethiopia

n Morocco,
w & c Europe to
Finland, Ukraine, | /
Kryra, Georgia,
ne Turkev

sw Saudi Arabia

n Cape Prov.
to e Zimbabwe,
c Mozambique

Angola (Huila)

e Zaire, s Tanzania
e Zambia, n Malawi

e Uganda , i
Kenya ,

n Tanzania C

+J Tunisia, e Algeri

East Africa

Fig. 13 Sects 27. Adenosepalum and 28. Elodes. Distribution of the 25 species. For key to annotations see Fig. 3 (p. 00) except for minor disjunctions,
which are indicated in two degrees, lesser ( ) and greater ( — ).

intermedium occurs in the Derna massif of NE Libya (24. H.
decaisneanum), having presumably reached that area when the
Saharan region had an equable climate.

A more extreme disjunction, however, separates H. annulatum
subsp. intermedium from 18b. subsp. annulatum, a plant of the
Macedonian region of the Balkan Peninsula and also two mountains
in Sardinia (Arrigoni et al., 1973). Despite the wide separation of
these populations from each other and the even wider separation of
both from the nearest African population, the morphological varia-
tion is such that the European populations together only just merit
subspecific rank. Despite this relative lack of variation, there is no
evidence that they reached their present area by long-distance
dispersal; and the occurrence of related derivative species in the
Aegean and eastern Mediterranean regions also favours the theory of
an ancient land-migration of H. annulatum.

These relatives of H. annulatum form two morphoclines. Two
relatively longer-haired species occupy isolated areas in the west
and north Aegean: the erect 19. H. delphicum in the islands Evvoia
(Euboea) and Andhros and the spreading 20. H. athoum in the Athos

Peninsula of northern Greece and the northern islands of Thasos and
Samothraki. These are morphologically quite distinct from H.
annulatum, but the other, south-eastern morphocline is not so well
isolated. 2 1 . H. atomarium occurs on both sides of the Aegean, in the
Peleponnisos and western Turkey, as well as on the eastern islands of
Lesvos, Khios, Ikarfa and Rhodos. In Khios and Ikaria a reduced
'form' of it, which is found in higher and more exposed habitats, is
treated here as a species, 22. H. cuisinii, although the morphological
'gap' between it and island plants of H. atomarium appears to be
small and could conceivably prove to be non-existent. A more
reduced form of H. cuisinii is found on Karpathos and apparently
Khasos, and the typical form also occurs on the Turkish mainland
(Boz Da£). The morphocline continues eastward in 23. H.
lanuginosum, of which the distribution slightly overlaps that of H.
atomarium in the Antalya region of south-western Turkey. From
there the area extends round the Mediterranean margin as far as
Israel (Judaea) and Jordan (Gilead), and this species also occurs in
northern Cyprus. A specimen from the 1830s labelled 'Sinai' indi-
cates, if the label is to be believed, that H. lanuginosum was once

92

N.K.B. ROBSON

found further south; but I know of no more recent records from there. Bermuda, Azores (introduced?) 29 species (+ 2 subspecies).

MorphologicallyextremepopulationsinsouthernTurkeyfromhigher 2Q M iandm (excluding garden

altitudes ('van scabrellum) and the extreme west of vil. Antalya . ^ . , ,

( ' var. pestalozzae ) are linked to the more typical form by a continu- ^

ous series of intermediates. ' SePals and Petals each 5 (excePl in occasional n°wers in SPP- l > 3'

22) or, if 4(3), then sepals small, subequal, not enclosing capsule
2

~ — ' ! Sepals and petals each 4; sepals in markedly unequal pairs, the outer

SYSTEMATIC TREATMENT4 iarge, enclosing capsule (subsect. 5. Ascynm) 27

2(1) Leaves articulated at base, sessile or pseudopetiolate, not

Sect. 20. MYRIANDRA (Spach) R. Keller in Engl. & amplexicaul; stamens deciduous (subsect. 1. Centrosperma) 3

Prantl, Nat. Pflanzenfam. 3(6): 214 (1893). Leaves not articulated at base or, if so (Sp. 23), then amplexicaul;

stamens persistent (except in Spp. 16, 17,23) 18

Shrubs, shrublets or perennial herbs up to 4.5 m tall, (the shrubs)

deciduous, glabrous, without dark glands; branching dichasial/ 3(2) Leaves with lamina expanded when mature, visible on both sides of

monochasial or rarely pseudo-dichotomous or mixed. Stems 2-4(- midrib; sePals exPanded 4

6)-lined and usually compressed (ancipitous) when young, usually Leaves acicular even when mature, usually ± only midrib visible

becoming terete, apparently eglandular; cortex exfoliating in flakes beneath; sepals acicular 9

or strips; bark thin and smooth or corky and sometimes with ^ Inflorescence ,_3(_7 or rarely more).flowered from uppermost

laticifers. Leaves opposite, decussate, sessile to subsessile, free, node sometimes with single flowers at 1(2) nodes below 5

deciduous at basal articulation (where present) or persistent; lamina

entire, with venation pinnate and closed to 1 -nerved, with tertiary Inflorescence (l)3-c. 25-flowered from uppermost node, with tri-

venation densely to laxly reticulate or absent; laminar glands punc- ads' dichasia or ""^"8 branches from l~

tiform; marginal gland dots dense; ventral glands absent. 5(4) Gynoecium 3-merous; leaves 8-22 mm wide, margin plane or

Inflorescence 1 -oo(over 70)-flowered from uppermost node, some- subrecurved; plant (0.6-) 1-3 m tall; sepals in flower 4-10 mm wide

times also from up to 10 nodes below, branching dichasial or rarely 1- frondosum

pseudo-dichotomous or mixed, sometimes with subsidiary flower- Gynoecium usually 5-merous; leaves 3-7(-10) mm wide; margin

ing branches from up to 8 lower nodes; bracts and bracteoles usually subrecurved to revolute; plant 0.14-0.6(-1 ) m tall; sepals in flower

reduced, rarely transitional. Flowers stellate, homostylous. Sepals 1.5-5 mm wide 3. kalmianum

(3-)4-5(-6), free, persistent and spreading to reflexed or deciduous

. . . , ,, . . , , .- 6(4) Inflorescence broadly to (usually) narrowly elongate-cylmdnc with

in fruit, with margin entire; veins l-5(-9); laminar glands puncti- 1.5.^** lateral dichasia from 2^ nodes; capsule 7-13 mm

form; marginal glands rather sparse, submargmal and mframargmal ,ong or stems 6 ]ined when young 7

glands absent. Petals (3-)4-5(-6), deciduous, with apiculus present,

lateral or rarely subapical, apiculate to obtuse or acute, or absent; Inflorescence shortly and broadly globose-cylindric or obpyramidal

margin entire, marginal glands absent; laminar glands sparse, with (2-)5-15-flowered lateral dichasia from 1-3 nodes; capsule 5-

..r .- , . c f • , A <r t .- t- 7 mm long; stems 4-lmed when young 8

stniform to punctiform, or absent. Stamen fascicles 4-5, not distin-
guishable individually, forming continuous narrow to broad band of 7(6) Leaves 30-70 mm long, narrowly oblong to narrowly elliptic or
30-650 stamens (i.e. 6-30 per fascicle), stamens individually de- oblanceolate; flowers 15-30 mm in diam.; placentation incom-
ciduous or persistent; filaments free; anthers yellow, gland amber; Plete'y axile; caPsule 7~13 mm lon§; seeds l-5~2 "jjj^jf
pollen type Vn. Ovary with 2-5 incompletely axile to parietal

placentae, each oo-ovulate; styles 2-5(6), free but mutually wholly Leaves 15-32(-37) mm long, very narrowly oblong-elliptic or

or partly appressed at anthesis, later often ± spreading or recurved; oblanceolate to linear; flowers 9-14 mm in diam.; placentation

stigmas minute. Capsule 2-5-valved, subcoriaceous, not vittate. parietal; capsule 4.5-6 mm long; seeds 0.7-0.8 mm long

Seeds ± broadly cylindric, carinate or not, without apical expansion; 6- 8alloldes

testa linear-reticulate or reticulate to linear-foveolate or scalariform- g^) Ovary (3-)4-5-merous; capsule deeply lobed; sepals narrowly el-
reticulate or scalariform. liptic or narrowly oblong to oblanceolate-spathulate, basal veins

3_7 4. lobocarpum

BASIC CHROMOSOME NUMBER (x). 9, 8; ploidy 2, 3 (cf. Sp. 5), 4?

(cf. Sp. 1). Ovary 3-4(-5)-merous; capsule not or scarcely lobed; sepals nar-
rowly oblong-lanceolate or narrowly oblong to oblong-spathulate,

HABITAT. Open, dry (cedar glades, barrens, shales, river bottoms, basal yeins ,_3 5 densinorum

rocky slopes, dry woods) or damp to wet (moist crevices, ditches,

seepage areas, meadows, marshes, bogs, swamps, ponds and 9<3) Stems 6-'ined when y™^ usual'y decumbent and mat-forming

lakesides), on granite or limestone or (Cuba) white sand; 0-1560 m

(N. America), -1650 m (Jamaica), -1800 m (Guatemala), -2000 m Stems 4-lined when young, erect 1 1

(Cuba), -2900 m (Hispaniola). ]Q(9) Leaves 4-1 1 mm long when mature; flowers sessile or almost so;

DISTRIBUTION. Canada (Newfoundland to Ontario), eastern U.S.A. sepals 1-4 mm long 7. tenuifolium

(westward to Minnesota, Iowa, Kansas, Oklahoma and Texas), east- Leaves 13-25 mm long when mature; flowers with pedicels c. 0.5

ern Mexico, Belize, Guatemala, Honduras, Greater Antilles, Bahamas, mm; sepals (3-)4.5-7 mm long 8. lloydii

1 1(9) Capsule cylindric or rarely very narrowly conic or ovoid-conic; leaf

4 In this part (Part 6), an asterisk (*) before a locality or after a herbarium symbol midrib below level of inrolled margins; bark smooth but not metal-
indicates that the specimen has not been seen by me. After a type specimen in sect. 20 lic-silverv 1 2
Mvriandra, it indicates that that type has been seen by Dr Adams. As in previous parts,
type material seen by me is indicated by an exclamation mark (!). Capsule ± narrowly ovoid-conic to ovoid or ellipsoid; leaf midrib

STUDIES IN HYPERICUM

93

level with inrolled margins but separated from each by a papillae-
lined groove; bark smooth and metallic-silvery or corky to spongy
16

12(1 1) Sepals articulated, deciduous; inflorescence overall mostly cylin-
dric, from up to 7 nodes, branching dichasial 13

Sepals not articulated, persistent; inflorescence overall obconic,
from terminal node only, branching mixed dichasial/pseudo-di-
chotomous 10. limosum

13(12) Capsule (4.5-)5-7 mm long; styles shorter than ovary; mature
leaves mostly 1 0—26 mm long, equalling or slightly exceeding those
in axillary clusters (9. nitidum) 14

Capsule 3.5-5 mm long; styles longer than ovary; mature leaves
mostly 6-1 1 mm long, usually twice as long as those in axillary
clusters 1 1. brachyphyllum

14(13) Leaves coriaceous with apex rounded to rounded-apiculate; petals
mostly 8-10 x 5-6.5 mm; sepals rounded-apiculate to obtuse
9a. nitidum subsp. cubense

Leaves subcoriaceous to chartaceous with apex rounded-apiculate
to long-acuminate; petals mostly 5-7 x 3-4 mm; sepals shortly
apiculate to acute 15

15(14) Leaf apex rounded-apiculate to obtuse, margin loosely inrolled;
sepals shortly apiculate to acute; plant to 3 m or more tall with ±
numerous stout bushy-branched stems from base
9b. nitidum subsp. nitidum

Leaf apex acute to long-acuminate, margin tightly inrolled; sepals
acute to long-acuminate; plant to c. 1 m tall with few slender little-
branched stems from base 9c. nitidum subsp. exile

1 6( 1 1 ) Bark smooth, metallic-silvery, without laticifers, exfoliating in large
thin curled plates; stylesc. 5 mm long; seeds 1-1. 6 mm long; young
stems, leaves and sepals strongly glaucous 12. lissophloeus

Bark thin corky to thick corky and spongy, containing ± conspicu-
ous vertical laticifers, exfoliating in thin papery sheets or plates;
styles 2.5^4- mm long; seeds 0.4-0.8 mm long; young stems, leaves
and sepals not glaucous 17

17(16) Stem internodes persistently ancipitous-winged when young; bark
with cork layers thin, smooth and containing inconspicuous thread-
like laticifers, not becoming thick on old stems; terminal
inflorescence (3-)7-32-flowered 13. fasciculatum

Stem internodes soon terete when young; bark with cork layers 3-4
mm thick, striate on account of coarse laticifers, becoming thick and
spongy on old stems; terminal inflorescence 1-3-flowered
14. chapmanii

18(2) Plant a decumbent mat-forming broad-leaved dwarf wiry shrub;
inflorescence 1-3-flowered (subsect. 2. Pseudobrathydium)
15. buckleyi

Plant an erect shrub or decumbent to rhizomatous subshrub or herb,
if broad-leaved then a herb or inflorescence °°-flowered 19

19(18) Sepals very unequal to subequal, persistent; stamens 120-c. 200;
inflorescence widely branched (subsect. 4. Brathydium) 20

Sepals unequal and deciduous or subequal to equal and persistent;
stamens 30-95; inflorescence narrowly branched (subsect. 3.
Suturosperma) 21

20(19) Leaves deciduous, (5-)7-20 mm wide, oblong-ovate or ovate to
triangular-lanceolate, base ± cordate-amplexicaul; erect (often
unbranched) shrub with bark on older stems corky
23. myrtifolium

Leaves persistent, 3-5 mm wide, linear-elliptic to linear, base nar-
rowly cuneate to rounded; decumbent ± branching subshrub with
bark thin or absent ... 24. dolabriforme

21(19) Sepals and usually stamens ± tardily deciduous; leaves oblong to
linear-oblong or elliptic to ovate-lanceolate, (7-) 10-25 mm wide;
placentation incompletely axile; shrubs 22

Sepals and stamens persistent; leaves mostly narrowly oblong or
triangular-oblong to linear, 1-10(-15) mm wide; placentation pari-
etal; shrubs, subshrubs or perennial herbs 23

22(2 1 ) Inflorescence terminal only, ( 1 )3-5(-8)-flowered; sepals 1 .5-3 mm
wide; capsules 6-14 x 4.5-7 mm; seeds with low ridge
16. apocynifolium

Inflorescence sometimes from more than one node, the terminal one
7-c. 45-flowered; sepals 1-1 .5 mm wide; capsule 3.5-7 x 3-5 mm;
seeds markedly carinate 17. nudiflorum

23(21) Inflorescence 1-3-flowered; perianth mostly 4-merous; leaves 1-3
mm wide; shrub 19. microsepalum

Inflorescence 7- 10- flowered or, if flowers 1-6, then plant herba-
ceous; perianth almost always 5-merous; leaves ( l-)3-l 5 mm wide;
shrubs to herbs 24

24(23) Capsule ovoid-cylindric to broadly ovoid; plant a shrub c. 0.5-1 .3 m
tall, base unbranched 18. cistifolium

Capsule broadly ovoid to depressed-globose or, if ovoid-ellipsoid to
ellipsoid, then plant a rhizomatous herb to 0.8 m tall and branched
from base 25

25(24) Plant a subshrub, not or rarely rhizomatous; capsule broadly
ovoid to depressed-globose; seeds 2-2.7 mm long
20. sphaerocarpum

Plant a rhizomatous herb; capsule ovoid-ellipsoid; seeds 0.6-0.7
mm long 26

26(25) Leaves narrowly oblong or narrowly lanceolate to linear
(l:b = c. 6-8); plant relatively stout, usually 0.4-0.8 m tall
21. adpressum

Leaves elliptic or ovoid-elliptic to oblanceolate (l:b = 2-4); plant
relatively slender, usually 0.1-0.2 m tall 22. ellipticum

27(1) Styles and placentas 3(4); stamens 70-100 28

Styles and placentas 2; stamens 30-50 30

28(27) Leaf base cordate-amplexicaul; inflorescence terminal, branching
pseudo-dichotomous 26. tetrapetalum

Leaf base cuneate to truncate or very rarely subcordate-amplexicaul;
inflorescence from 1-5 nodes, branching dichasial and/or pseudo-
dichotomous 29

29(28) Outer sepals apiculate to rounded; leaves without basal gland-like
auricles, margin plane to subrecurved; terminal inflorescence l-3(-
7)-flowered, branching nearly always wholly dichasial
25. crux-andreae

Outer sepals acute to subacuminate; leaves with basal gland-like
auricles, margin subrecurved to subincrassate; inflorescence 1-
flowered, branching pseudo-dichotomous 27. edisonianum

30(27) Pedicels elongate (bracteoles near uppermost leaves), reflexed soon
after anthesis; leaves without basal gland-like auricles
28. suffruticosum

Pedicels short (bracteoles close to sepals), persistently erect; leaves
with basal gland-like auricles (29. hypericoides) 31

31(30) Plant erect; stems 0.3-1.5 m, unbranched from base but freely
branched above; leaves oblong to linear, 7-25 mm long
29a. hypericoides subsp. hypericoides

Plant decumbent to prostrate; stems 0.05-0.3 m, branched from
base; leaves oblanceolate or oblong-spathulate or, if narrowly ob-
long, then 3-10 mm long 32

94

N.K.B. ROBSON

32(31) Plant decumbent; leaves usually oblanceolate, 10-20 x 3-6 mm

long; inflorescence-branching dichasial or lateral

29b. hypericoides subsp. multicaule

Plant prostrate; leaves narrowly oblong to oblong-spathulate, 3-8(-
10) x 1-2.5 mm long; inflorescence-branching pseudo-dichotomous
29c. hypericoides subsp. prostratum

Subsect. 1. Centrosperma R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 3(6): 214 (1893). Type: H. prolificum L. (lectotype,
W.P. Adams, 1962).

Shrubs with leaves articulated at base, deciduous; inflorescence-
branching dichasial (or mixed dichasial/pseudo-dichotomous in 10.
H. limosum), mainly basipetal; sepals 5 (very rarely 4), very unequal
to subequal or rarely equal, deciduous; petals 5 (rarely 4); stamens
30-650, deciduous; styles and placentae (2)3-5(6), placentation
incompletely axile to parietal. Species 1-14.

Hypericum frondosum Michx., Fl. bor.-amer. 2: 81 (1803);
Poir., Encycl., Suppl. 3: 694, 699 (1813); Pursh, Fl. Amer. sept.:
375 (1814); Choisy, Prodr. monogr. Hyperic:. 38 (1821), in DC.,
Prodr. 1: 554 (1824); Spach in Annls Sci. nat. (Bot.) II, 5: 364
(1836) sub Roscyna sine comb.; Sprague in Curtis' s hot. Mag.
139: t. 8498 (1913); Lott in J. Arnold Arbor. 19: 149 (1938);
Svenson in Rhodora 42: 15 (1940); Rehd., Man. cult, trees 2nd
ed.: 640 ( 1 940); W.P. Adams in Contr. Gray Herb. Harv. no. 1 89:
16(1 962), in/ Elisha Mitchell sclent. Soc. 89: 69 ( 1 973); Correll
& Johnston, Man. Vase. PL Texas: 1063 (1970); R.C. Clark in

Map 1 Sect. 20: 1. H. frondosum • specimens, O records.

Ann. Mo. hot. Gdn 58: 209 ( 1 97 1 ); Bean, Trees & shrubs hardy in
Br. Isles 8th ed. 2: 413 (1973); Clewell, Guide vase. pi. Florida
Panhandle: 372 ( 1 985); Godfrey, Trees, shrubs & woody vines N.
Florida, etc. : 370 ( 1 988); N. Robson in Cullen et al., Eur. Gdn Fl.
4:67,ff. 10.1, 1 0.9 (1995). Type: U.S.A., Tennessee?, 'ad flumen
Tennassee', n.d. (st), Michauxs.n. (P-holotype, BM!-microfiche;
GH*-photograph & sketch).
Fig. 14A, Map 1.

H. aureum W. Bartram, Travels Carolina: 383 (1791); Torrey &
Gray, Fl. N. Amer. 1: 161 (1838); Chapm., Fl. South. U.S.: 40
( 1865); Coulter in Bot. Gaz. 11: 84(1886), in A. Gray, Syn.fl. N.
Amer. 1: 286(1897); Small, Fl. s.e. U.S.: 790 (1903), Man. s.e.fl.:
872 (1933); Sprague in Curtis 's Bot. Mag.: t. 8498 (1913); R.A.
Vines, Trees, shrubs & woody vines of S.W.: 754 (1960); N.
Robson in Cullen et al., Eur. Gdn Fl. 4: 67, ft". 10.1, 10.9 (1995),
non Loureiro ( 1 790). Type: U.S.A., Georgia, Flint R. Taylor Co.,
Patse-Liga [Patsilaga] creek, 1776 (fl), Bartram Book D/5 (BM!-
holotype).

H. amoenum Pursh, Fl. Amer. sept. 2: 375 (1814). Type: U.S.A., 'in
South Carolina and Georgia', Lyon s.n. (OXF-holotype).

H. rugelianum Kunze in Linnaea 24: 1 77 ( 1 85 1 ). Type: cultivated in
Leipzig Bot. Card, from seed ex U.S.A., Tennessee, Oberland
Mts, Rugel (LZt-holotype).

Brathydium aureum (W. Bartram) K. Koch, Hon. dendroi: 66
(1853).

B. rugelianum (Kunze) K. Koch, Hon. dendroi:. 67 (1853).

Hypericum prolificum [var.] P aureum (W. Bartram) Koehne, Deut.
Dendroi:. 416 (1893).

H.splendensSmattmBull.Torr.bot. Club29:29\ (\90\),Fl.s.e. U.S.:
790 (1903), Man. s.e.fl.: 872 (1933); Svenson in Rhodora 42: 15
(1940). Type: U.S.A., Georgia, De Kalb Co., Stone Mt., 506 m, 4
July 1893 (fl), Small s.n. (NY-holotype; A!, F!, US!-isotypes).

Icon: Sprague in Curtis' s hot. Mag. 139: t. 8498 (1913).

Shrub (0.6-)l-3 m tall, erect, much branched above base, with
branches lateral, strict to spreading, forming rounded bush or small
'tree'. Stems green, 4-lined and ancipitious when young, soon 4-
lined and rounded, becoming reddish brown and 2-lined to terete in
2nd or 3rd year; cortex exfoliating in strips or (often large) plates;
bark pale grey, smooth, thin. Leaves sessile, ascending to spreading;
lamina 25-65 x 8-22 mm, oblong to lanceolate-oblong or some-
times narrowly elliptic to oblanceolate, plane or with margin
subrecurved, paler or somewhat glaucous beneath or wholly ±
glaucous, chartaceous, deciduous at basal articulation, apex apiculate-
obtuse to rounded, base broadly cuneate and attenuate to narrowly
cuneate; venation sometimes rather obscure: c. 10-16 pairs main
laterals with subsidiaries and densely reticulate tertiaries, midrib
prominent; laminar glands dense. Inflorescence l-3(-7)-flowered,
sometimes with paired single flowers or triads or 1-3-flowered
branches at node below and very rarely single flowers at third node
('//. splendens'); pedicels 1.5 (terminal)-lO mm long; bracts foliar
or reduced, oblong-elliptic. Flowers 25-45 mm in diam.; buds
globose. Sepals 5(4), 6-14(-20) x 4-10 mm, enlarging in fruit,
imbricate, very unequal, ovate or oblong to elliptic or elliptic-
spathulate or sometimes foliaceous, rounded or apiculate-obtuse,
plane or margin recurved, basal veins (3)5-7(9), branching and
densely reticulate distally. Petals 5(4), golden yellow to orange-
yellow, base soon brownish, becoming incurved-deflexed, 1 2-25 x
6-14 mm, 1.25-2 x sepals, obovate to oblanceolate, with apiculus
lateral (petals sometimes bifid), rounded. Stamens c. 250-650,
longest 9-12 mm, 0.45-0.75 x petals. Ovary 3-merous, 6-8 x 3-4(-
5) mm, ± narrowly pyramidal-ovoid to broadly ellipsoid, acute to
apiculate-obtuse, placentation incompletely axile; styles 3, 4-6 mm
long, 0.65-0.75 x ovary, remaining erect. Capsule 1 2-1 5 x 6-8 mm,
narrowly ovoid-conic to broadly ovoid-rostrate, rounded-trigonous,
exceeding or shorter than sepals and often initially surrounded by
them, thickly coriaceous. Seeds blackish brown, c. 1 .5 mm long,
carinate; testa shallowly linear-reticulate. 2n = 18 (Hoar & Haertl,
1932; Adams in Robson & Adams, 1968), 36 (see note below).

STUDIES IN HYPERICUM

95

Fig. 14 A. H.frondosum: (a) habit; (b) leaf (part); (c) sepal; (d) petal; (e) stamen; (0 anther; (g) ovary; (h) ovary, T.S.; (i) capsule. B. H. kalmianum: (j)
habit; (k) capsule. C. H. prolificum: (1) habit; (m) capsule (a, j, 1 x Vr, b-d, g, i, k, m x 4; f x 6). A. Krai 39725. B. Bar/7 2256. C. Demaree 45955.

96

N.K.B. ROBSON

Dry cedar glades and barrens on limestone and calcareous shales,
also (Demaree 405749) 'swampy creek bottoms'; 180-506 m.

U.S.A. (southwestern end of the Appalachian Range), in northern
and central Georgia, northern Alabama, eastern Mississippi, north-
ern Louisiana, eastern Tennessee and southern Kentucky; also
recorded from eastern Texas. Adventive in Connecticut, New York
and Massachusetts (fide Adams, 1962).

U.S.A. Alabama: FranklinCo.,Russelville,26June 1970(fl),Kra/39725
(BM); *Morgan Co., 3.2 km S. of Tennessee R., S. of Huntsville, Godfrey
57520 (FSU); Lawrence Co., 1.75 km S. of Moulton, 26 June 1970(fr), Krai
41278(BM). Georgia: DeKalbCo., Stone Mt,NW slope, 390m, lOJuly 1900
(fl). Wilson 13 (BM, K); Quitman Co., Chattahoochee R. above Georgetown,
16 October 1902 (fr). Harper 1755 (BM). Kentucky: *Crittenden Co.,
Dycusburg overlooking Cumberland R., 16 June 1969 (fl), Athey 695 (MO);
* Wayne Co., S. of Monticello, Smith & Hodgdon 3978 (GH, NY). Louisiana:
*Natchitoches Par., c. 6.4 km E. of Provencal, Correll 9829 (DUKE, NY).
Mississippi: *ClarkCo.,6October 1929,/U/H>s.n.(UNC).Tennessee: Bedford
Co., 4 km S. of Shelby ville, 29 August 1 958 (fr), Godfrey 57535 (BM, FSU*);
Rutherford Co., La Vergne, 180m, 24 June 1962 (fl), Demaree 45749 (BM);
Wilson Co., CedarGlades, 3.2 km N. of Lebanon, 28August 1 958 (fr), Godfrey
57498 (BM, FSU*). Texas: fide Correll & Johnston (1970).

CULTIVATED. Specimens seen from England (1904-1988), Holland
(1920) and U.S.A. (1897-1959).

H.frondosum has the most primitive characters of any species in sect.
Myriartdra, its nearest (ancestral) species being H. synstylum N.
Robson from Ethiopia and Somalia (sect. 1 . Campylosporus). Only
one character, trimery of the gy noecium, might be regarded as special-
ized in comparison with 3. H. kalmianum L. and 4. H. lobocarpum
Gatt. Since these species, however, are much more specialized in their
other characters, there are grounds for assuming that pentamery of the
gynoecium is advanced, not primitive, in this section; and its constant
trimery in H.frondosum would support this assumption.

H.frondosum is the nodal species in sect.Myriandra, being related
to species in three subsections. Through 2. H. prolificum it is related to
the rest of subsect. Centrosperma, through 23. H. myrtifolium to
subsect. Brathydium, and through 25. H. crux-andreae to subsect.
Ascyrum. From//, prolificum it can be distinguished (at least in natural
habitats) by the larger leaves, flowers and fruits and by the virtual
restriction of the inflorescence to the terminal node. When flowers are
present at lower (1-2) nodes, they are solitary or rarely 3 in each leaf
axil. The larger floral parts also help to distinguish//, frondosum from
the other two related species. In addition, H. myrtifolium has cordate-
amplexicaul leaves, whilst H. crux-andreaehasatetramerous perianth
and spreading styles. At least in cultivation, H. frondosum itself
produces a few flowers with a tetramerous perianth, thus indicating
where 'Ascyrum ' has been derived from Hypericum.

The flowers of H. frondosum rarely exceed 45 mm in diameter,
and the diploid chromosome number is normally 1 8. One specimen
in the Arnold Arboretum herbarium (A), however, has a flower 60
mm in diameter and is labelled as a tetraploid (2n = 36). It was
collected from the Arboretum Trial Ground (J. W. Peterson J- 1 96).
Perhaps the extraordinarily large-flowered cv. Sunburst is also tetra-
ploid. Colchicine-induced tetraploids of this species have been
produced (Myers, 1963).

Ix. Hypericum frondosum x prolificum

H. x vanfleetii Rehd., Man. cult, trees: 640 (1940) ['Van Fleetii'];
W.P. Adams \nRhodora 74: 281 (1972). Type: cultivated 1925 (at
Arnold Arboretum, Jamaica Plain, Mass.?), not seen.

'Flowers 2.5-3 cm across, in terminal cymes, often with a few pairs
of axillary solitary fl[owe]rs below' [Render].

Although the above species appear to remain distinct in the field,

they hybridize in cultivation; and artificial hybrids between them
have been made (Myers, 1963). There is a series of garden forms,
intermediate in size of parts between H.frondosum and H. prolificum
and with intermediate inflorescence forms, that breaks down the
only differentiating character other than size. According to Adams,
Rehder's description of H. x vanfleetii would apply to pure H.
frondosum, whereas all specimens labelled H. x vanfleetii in the
Arnold Arboretum Herbarium could easily be accommodated in H.
prolificum. The use of the term 'cymes' by Rehder, however, sug-
gests that these comprise more numerous flowers than is normal in
H. frondosum; and the virtual restriction of the inflorescence to the
terminal node is atypical of H. prolificum. One of Rehder's speci-
mens, in my opinion, agrees with his description (C.E.K. & C.K.A.
Arbor. 20895), but the other two, as Adams has said, are probably
pure H. prolificum. None of these specimens is the type, as none has
the date given by Rehder. In the absence of a definite type, or until
one such is discovered, I therefore suggest that it is appropriate to
use the specific epithet vanfleetii for those cultivated intermediates
(and any wild ones that may occur), in the supposition that they are
of hybrid origin (see also other Rehder hybrid names, pp. 1 00, 101).

2. Hypericum prolificum L., Mant. pi. 1: 106 (1767); Lam., Tab.
encycl 2: t. 643 f. 2 (1796), Encycl. 4: 159 (1797); Pursh, Fl
Amer. sept. 2: 375 (1814); Choisy, Prodr. monogr. Hyperic.: 46
(1821)proparteexcl. syn.//. kalmianum L., in DC., Prodr. 1: 547
( 1 824); Torrey & Gray, Fl. N. Amer. 1 : 1 59 ( 1 838) pro parte excl.
syn. H. densiflorum; S. Watson, Bibliogr. index N. Amer. hot. 1:
128 (1878); Coulter in Bot. Gaz. 11: 84 (1886), in A. Gray, Syn.
fl. N. Amer. 1: 285 (1897); Sargent in Gdn Forest 3: 525, f. 66
( 1 890); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21:
180 (1925); Svenson in Rhodora 42: 9 (1940); Rehd., Man. cult,
trees, 2nd ed. 640 (1940); Fernald & Schubert in Rhodora 50:
167, t. 1 101 ff. 1-3 (1948); Svenson \v\Rhodora 54: 205 (1952);
W.P. Adams in Rhodora 61: 250 (1959), in Contr. Gray Herb.
Harv. no. 189: 15 (1962), in J. Elisha Mitchell scient. Soc. 89: 69
(1973); R.A. Vines, Trees, shrubs & woody vines of S.W.: 755
(1960); Radford, Ahles & Bell, Man. vase. fl. Carolinas: 712
(1968); Utech & Iltis in Trans. Wis. Acad. Sci. Arts Lett. 58: 335
(1970); R.C. Clark in Ann. Mo. hot. Gdn 58: 209 (1971); Bean,
Trees & shrubs hardy in Br. Isles 8th ed. 2: 422 (1973); J.M.
Gillett & Robson in Puhls Bot. natn. Mus. nat. Sci. Can. no. 11:
9, t. 3, map 2 ( 1 98 1 ); Cooperrider in Castanea 54: 7, f. 1 ( 1 989);
N. Robson in Cullen et al., Eur. Gdn FL 4: 67, ff. 10.1, 10.10
(1995). Type: U.S.A., without precise locality, Herb. Linn. 943/
20 (LINNI-lectotype, Svenson, 1940). For discussions of
Svenson's choice, see Fernald & Schubert (1948), Svenson
(1952), Adams (1959, 1962) and Gillett & Robson (1981).

Fig. 14C, Map 2.

H. cryptopetalumVogel inTrew, PL rar., dec. 3: 1, t. 21 f. 1 (1784),
nom. illegit. (Art. 63). Type as for H. prolificum L.

H.foliosum sensu Jacq., PL hort. schoenbr. 3: t. 299 (1798) pro parte
quoad descr. et tab., non Aiton (1789).

Myrian dra prolifica (L.) Spach, Hist. nat. veg. Phan. 5: 439 (1836),
inAnnlsSci. nat. (Bot.) II, 5: 365 (1836); K. Koch, Hort. dendrol.:
66(1853).

M. spathulata Spach \nAnnlsSci. nat. (Bot.) II, 5: 365 (1836). Type:
U.S.A., 'dans les provinces meridionales des Etats-Unis', Leconte
s.n. (P-holotype).

M. ledifolia Spach, Hist. nat. veg. Phan. 5: 441 (1 836), in Annls Sci.
nat. (Bot.) II, 5: 365 (1836). Type: U.S.A., Florida?, 'dans le midi
Etats-Unis', Leconte s.n. (P!-holotype). Coulter (1886a) treated
M. ledifolia as a synonym of//, prolificum, but, as Adams (1962:
50) pointed out, the description is not diagnostic; it could refer to

STUDIES IN HYPER1CUM

97

Map 2 Sect. 20: 2. H. prolificum • specimens, O records; 3. H.
kalmianum • specimens, D records.

H. densiflorum. According to my own records, however, the

specimen does belong to H. prolificum.
Hypericum spathulatum (Spach) Steud., Nomencl. 2nd ed. 1: 789

(1840); Fernald & Schubert in Rhodora 50: 168, t. 1101 f . 4

(1948); Fernald, Gray's Man. Bot. 8th ed.: 1011 (1950); J.P.

Gillespie in Castanea 24: 29 (1958); Mohlenbr., ///. Fl. Illinois fl.

pis, Hollies to Loasas: 37, t. 15 (1978).
Myriandra prolifica [var.] j spathulata (Spach) K. Koch, Hort.

dendrol.: 66 (\853).

Brathys prolifica (L.) Payer, Traiteorganogen.fi, 8: t. 1 (1857).
Hypericum kalmianum var. majus Gatt. in Bot. Gaz. 11: 275 (1886),

Tennessee FL: 29 (1887), nomen.
H. prolificum var. montanum Gatt., Tennessee Fl. : 29 ( 1 887), nomen.

Icones: Sargent in Gdn Forest 3: 524, f. 66 ( 1 890); Jacquin, PL hort.
schoenbr.3:t.299(179&).

Shrub (0.2-)0.75-1 .5(-2) m tall, erect (or ± diffusely branched) with
branches erect to ascending, forming rounded or irregular bush. Stems
green, 4-lined and ancipitous when young, soon 4-lined and rounded,
becoming reddish brown and terete in 2nd or 3rd season; cortex
exfoliating in strips or flakes; bark pale grey, smooth, thin. Leaves
sessile or rarely with pseudopetiole to 3(4-6) mm long, sometimes
clustered; lamina 30-70 x 6-15 mm, narrowly oblong to narrowly
elliptic-oblanceolate, plane? or with margin ± recurved, paler or
somewhat glaucous beneath, deciduous at basal articulation, apex

rounded-apiculate or rarely retuse to acute, base attenuate to narrowly
cuneate (and then pseudopetiolate); venation usually clearly visible
beneath: c. 10-16 pairs main laterals with subsidiaries and densely
reticulate tertiaries, only midrib prominent; laminar glands dense. In-
florescence ( 1 )3-7(-9)-flowered, rarely with paired accessory flowers,
with paired single flowers or triads or 1 -3(-7)-flowered branches from
2 nodes below, the whole broadly to narrowly cylindric; pedicels 1-2
mm long; bracts foliar or reduced, oblong-elliptic. Flowers 1 5-30 mm
in diam.; buds broadly ovoid. Sepals 5, 4-8 x 1 .5-^4- mm, enlarging in
fruit, imbricate, unequal to subequal, broadly or narrowly elliptic to
obovate-spathulate or oblanceolate, obtuse or apiculate-obtuse to
acute, plane or with margin recurved, basal veins 3-7, not or obscurely
branching. Petals 5, golden yellow, becoming incurved-deflexed, 7-15
x 3-6 mm, 1 .75-2 x sepals, obovate to oblanceolate-spathulate, with
apiculus lateral, obtuse or apiculate-obtuse to acute. Stamens c. 150-
500, longest 10-11 mm, 0.7-0.85 x petals. Ovary 3(4-5)-merous,
(3-)4— 5.5 x 1.2-3 mm, narrowly ovoid to narrowly ellipsoid, acute,
placentation incompletely axile; styles 3(4-5), 4-6 mm long, 1-1.3 x
ovary, remaining erect, separating slightly only as fruit matures.
Capsule (6-)7-l 3 x 4-7 mm, narrowly ovoid-conic to ovoid or rarely
ellipsoid, acute to subacute, trigonous to rounded, exceeding sepals,
thickly to thinly coriaceous. Seeds blackish brown, 1.5-2 mm long,
ecarinate; testa linear-reticulate to subscalariform. 2n = 1 8 (Nielsen,
1924; Adams in Robson & Adams, 1968).

Rocky slopes, embankments, dry river bottoms, open woodland (in
north), usually on limestone but sometimes on granite; 90-over
600m.

Eastern U.S.A., Canada (southern Ontario): north to Iowa, Michigan
and New York, west to Missouri, Arkansas and Oklahoma, south to
Mississippi, Alabama and Georgia.

CANADA. Ontario: near London (Gillett & Robson, 1 98 1 : map 2).

U.S.A. Alabama: *Madison Co., Huntsville, 1925, Newsom 8 (MO);
SumterCo.,S.ofDancy,byAla 17, 27 June 1 970 (fl & fr), /fra/ 39768 (BM).
Arkansas: Hot Spring Co., P.O. Malvern, Coastal Plain, Social Hill, 90 m, 4
July 1967 (fl), Demaree 56579 (BM); Montgomery Co., Ouachita R., South
Fork, 210 m, 1 October 1962 (fr), Demaree 46619 (BM). District of
Columbia: *High Island, 30 August 1905 (fr), House 1496 (MO). Georgia:
Gordon Co.?, Costanaula R. below Resaca, 192 m, December 1903 (fr),
Harper 2035 (MO); Heard Co., S. of Franklin, Camp Meeting Rock, 2
August 1958 (fl & fr), Adams 140 (K). Illinois: *Franklin Co., Big Muddy
River Bottoms, Plumfield, 22 July 1941 (fl & fr), McCree 939 (MO);
Johnson Co., Tunnel Hill, 15 May 1919 (fr). Palmer 15547 (MO). Indiana:
*Crawford Co., c. 1.6 km NE of Leavenworth, Deam 18585 (*A, *IND);
*Fulton Co., Tippecanoe R., 22 August 1926 (fl & fr), Churchill 573 (MO).
Iowa: *Des Moines Co., Burlington, 7 August 1925 (fr), Pammell65 (MO);
*Lee Co., 1.8 km W. of Donnellson, 15 July 1928 (fl), Shimek s.n. (MO).
Kentucky: *Bath Co., 5.6 km SW of Preston, 2 June 1938 (fr), Wharton 2474
(MO); ? Co., Kentucky R., Bronsborough, August 1934 (fl), Peter s.n. (K).
Louisiana: Natchitoches, 16 June 1915 (fl). Palmer 8008 (K). Maryland:
Cecil Co., Susquehanna River bottoms, 2km S. of Conowingo, ISJuly 1971
(fl), Lombardo & Windier 3622 (H); *Plymouth Co., Plymouth, 9 August
1884 (fl & fr). Gray s.n. (MO). Michigan: *St ClairCo., Port Huron, 21 July
1894 (fl), Dodge s.n. (MO); ? Co., Kalamazoo, n.d. (fl), Hb. Carey (K).
Mississippi: one record (fide Adams). Missouri: Carter Co., Van Buren, 4
July 1914 (fl?). Palmer 6195 (K); Jefferson Co., Mammoth Creek, 10.5 km
W. of De Soto, c. 170 m, 27 July 1895 (fl & fr), P. & T. Raven 16768 (BM,
MO). New Jersey: Jersey, n.d. (fl), Bartram s.n. (BM). New York: *Cortland
Co., Solon to Cincinnatus, 27 August 1916 (fr), Wiegand6Sl2 (MO). North
Carolina: McDowell Co., Curtis Creek road c. 3 km S. of Blue Ridge
Parkway, 25 July 1968 (fl), Leonard & Radford 1805 (BM, H); Richmond
Co., 7.2 km NW of Buckingham, 5 October 1956 (fr), Radford 19158 (H).
Ohio: Clermont Co., Loveland, 2 1 July 1 878 (fl), James 269 (K); *Lucas Co.,
NW of Whitehorse, 19 October 1919 (fr), Moseley s.n. (MO). Okla-
homa: Le Flore Co., W. end of Rich Mtn, Ouachita Nat. For., Page, 480 m, 23
October 1966 (fr), Demaree 53252 (BM); Logan Co., Strictland, 13 August
1937 (fr), Demaree 15752 (MO). Pennsylvania: Fayette Co., Ohio Pyle, 20

98

July 1902(fl),S/za/*r281 (BM); Northampton Co., Nazareth, 2 October 1849
(fr), Prior s.n. (K). South Carolina: Pickens Co., Table Rock, 1843 (fr), Gray
& Sullivant s.n. (K). Tennessee: Polk Co., Hwy. 30, Reliance, 15 July 1969
(fl), Rogers & Bowers 43918 (H); Maury Co., above Duck R. by 1-65, 18.7
km NNE of Lewisburg, 3 June 197 1 (fr), Krai 43539 (BM). Virginia: ? Co.,
Chain Bridge, 7 October 1928 (fr), Tanaka 7262 (TAI); *Prince George Co.,
Flowerdew Hundred, 23 July 1938 (fl & fr), Fernald & Long 8769 (MO);
Smyth Co., middle Holston Valley near Seven Mile Ford, 607 m, 1892? (fl),
Small s.n. (K). Wisconsin (naturalized, see Utech & Iltis, 1970: 335, map 2):
*Richland Co., 1 1.2 km W. of Boaz, 245 m, 6 August 1978 (fl & fr), Nee
16536 (MO).

CULTIVATED. Specimens seen from England (1842-1988), Ireland
(1968-1971), Holland (n.d.), Germany (1732-1827), Switzerland (s.n.),
France (1818-1890), Japan (1954-1955), China (1984), India (1956) and
U.S.A. (1922-1979).

H. prolificum is the pivotal species in subsect. Centrosperma, linking
1 . H.frondosum with 3.H. kalmianum and the//, densiflorum (Spp. 4-
8), H. nitidum (Spp. 9-11) and H.fasciculatum (Spp. 12-14) groups
respectively. H. kalmianum differs by the shorter habit and the
restriction of the inflorescence to the terminal node, and usually by the
4-5-merous gynoecium and narrower leaves; and the other groups
differ by the smaller flowers and (usually) fruits and usually narrower
leaves. Most of them also have less intrusive placentae.

H. prolificum is very variable, the most primitive forms occurring
in Arkansas and Oklahoma, i.e. on the other side of the Mississippi
valley from the major area of H.frondosum. Thence it has apparently
spread mainly northward and eastward round the north of the
Mississippi Embayment to Iowa, Wisconsin, Michigan, New York
and New Jersey and south to Mississippi, Alabama and Georgia. It is
rare in the eastern Coast Plain and absent from, although (according
to Adams, 1962: 15) hardy in, Massachusetts.

For hybrids with 1. H. frondosum, 3. H. kalmianum, 4. //.
lobocarpum and 5. H. densiflorum, see discussions under these
species respectively.

3. Hypericum kalmianum L.,Sp. pi.: 783 (1753); Lam.,Encycl. 4:
148 (1797); Pursh, Fl. Amer. sept. 2: 374 (1814); Choisy, Prodr.
monogr. Hyperic. : 4 1 ( 1 82 1 ) pro parte excl. syn. //. bartramianum
Mill., in DC., Prodr. 1: 545 (1824); Torrey & Gray, Fl. N. Amer.
1: 158 (1838); S. Watson, Bibliogr. index N. Amer. hot. 1: 127
( 1 878); Coulter in Bot. Gaz. 11: 83 ( 1 886), in A. Gray, Syn. Fl. N.
Amer. 1: 285 (1897); Sargent in Gdn Forest 1: 1 13, f. 24 (1890);
Sprague in Curtis' s bot. Mag. 139: t. 8491 (1913); R. Keller in
Engl. & Prantl, Nat. Pflanzenfam. 2nd ed 21: 180 (1925);
Svensonin/?/ioJ0ra42:9(1940);Rehd.,Mtfrt. cult. trees2nded.:
639 (1940); W.P. Adams in Contr. Gray Herb. Harv. no. 189: 12
(1962); Utech & Iltis in Trans. Acad. Sci. Arts Lett. 58: 329
(1970); Bean, Trees & shrubs hardy in Br. Isles 8th ed. 2: 416
(1973); Mohlenbr., ///. Fl. Illinois fl. pis, Hollies to Loasas: 29, f.
1 1 (1978); J.M. Gillett & Robson in Publs Bot. natn. Mus. nat.
Sci. Can. no. 11: 7, t. 2, map 2 (1981); Cooperrider in Castanea
54: 7, f. 1 (1989); N. Robson in Cullen et al., Eur. Gdn Fl. 4: 67
(1995). Type: U.S.A., 'habitat in Virginia' (probably New York,
near Niagara, see Gillett & Robson 1 98 1 : 9), Kalm in Herb. Linn.
943/2 (LINN-holotype; A, BM!-photographs).

Fig. 14B, Map 2.

Norysca kalmiana (L.) K. Koch, Hort. dendroi: 66 (1853).

Icones: Sprague in Curtis 's bot. Mag. 139: t. 849 1 ( 1 9 1 3); Mohlenbr.
///. Fl. Illinois fl. pis, Hollies to Loasas: 30, f. 1 1 (1978).

Shrub (0. 14-)0.2-0.6(-1) m tall, erect, with branches erect to
ascending, forming slender to rounded or flat-topped bush. Stem
green, 4-lined and ancipitous when young, soon 4-lined and
rounded, becoming reddish brown and terete in 2nd season; cortex
exfoliating in strips; bark smooth, thin. Leaves sessile, sometimes

N.K.B. ROBSON

in immature clusters in leaf axils; lamina (15-)20-45 x 3-7(-10)
mm, narrowly oblong to oblanceolate or linear, with margin
subrecurved to revolute, paler or ± glaucous beneath, chartaceous,
deciduous at basal articulation, apex rounded to obtuse or
subapiculate-obtuse, base narrowly cuneate to subattenuate; vena-
tion rather obscure beneath: c. 9-14 pairs main laterals with
subsidiaries and densely reticulate tertiaries, only midrib promi-
nent; laminar glands dense. Inflorescence ( 1 )3-7(rarely more)-
flowered, without accessory flowers, restricted to terminal node or
rarely also from 1-2 nodes below; pedicels 2.5-7 mm long; bracts
reduced, linear-oblong to oblanceolate. Flowers 20-35 mm in diam.;
buds broadly ovoid. Sepals 5(4), 4-9 x 1.5-5 mm, enlarging and
divergent to reflexed in fruit, imbricate, subequal to unequal (when
4), elliptic or oblong to obovate, obtuse or apiculate to acute,
margins recurved to revolute, basal veins 3-7, branching and reticu-
lating distally. Petals 5(4), golden yellow, becoming incurved-
deflexed, 8-15 x 5-9(-12) mm, 1.6-2 x sepals, obovate to oblong,
with apiculus lateral, rounded or obsolete. Stamens c. 150-200,
longest 6-10 mm, 0.65-0.75 x petals. Ovary (3-4)5(6)-merous, 4-
6 x 1.5-2 mm, narrowly ovoid, acute; placentation incompletely
axile; styles (3-4)5(6), 3-4 mm long, 0.65-0.75 x ovary, remaining
erect, separating only as fruit matures. Capsule 7-1 1 x 4-7 mm,
narrowly ovoid-conic to narrowly cylindric-ellipsoid, obtuse, rounded
or slightly lobed, longer than sepals, thinly coriaceous. Seeds pur-
plish brown, 0.7-1.1 mm long, shallowly carinate; testa
subscalariform. 2n = 18 (Hoar & Haertl, 1932; Pringle, 1976).

Dunes and sandy or calcareous rocky shores, sandy or calcareous
plains and low prairies, along rivers and in Sphagnum-sedge swamps;
c. 1 80-400 m.

U.S.A. and Canada adjacent to the Great Lakes and along the Ottawa
River. The record (van majus) for Tennessee (Coulter, 1886£;
Gattinger, 1887) is based on a mis-identification (see p. 97).

CANADA. Quebec: Pontiac Co., Plage Pontiac, 22 July 1942 (fl), Baril
2256 (BM); Pontiac Co., Bristol, 20 September 1938 (fr), Marshall s.n. (K).
Ontario: Ottawa Distr., Carleton Co., March Township, Ottawa River N. of
Shirley Bay, 14 August 1948 (fl), Colder & Cody 1583 (BM); Simcoe Co.,
Georgian Bay, Nottawasaga Bay, c. 1.6 km W. of Collingwood, 14 August
1974 (fl & e. fr), Gillett 16653 (H).

U.S.A. Illinois: Chicago, pre-1879 (fl), Vasey 182 (K); *Lake Co.,
Waukegan, Gleason & Shobe 331 (DUKE). Indiana: Lake Co., near Grand
Calumet, Miller, 1 July 1934 (fl), Buhl F669 (TAI); *Starke Co., Bass Lake,
Deam 201 13 (IND). Michigan: Mason Co., Ludington, 20 May 1956 (o. fr),
Rolland-Germain 6451 (K); Emmett Co., Wycamp Lake near Cross Village,
29 July 1933 (fl), Gleason & Gleason 261 (G, K). New York: Niagara Falls,
Table Rock, n.d. (fl), Gray s.n. (K). Ohio: near Toledo, ./We Adams (1959:
map 1). Wisconsin: Door Co., Mud Bay, 24 May 1968 (st), Borg s.n. (H);
*Juneau Co., N. of Mauston, 8 July 1936 (fl), Fassett 17928 (MO).

CULTIVATED. Specimens seen from England (1904-1967), Scotland
(1977), Germany (1910), Japan (1930), Australia (1889) and the U.S.A.
(1920-1974).

STUDIES IN HYPERICUM

99

Despite its 5-merous ovary, H. kalmianum is clearly a northern
derivative of H. prolificum, from which it differs in its shorter habit,
its inflorescence usually being confined to the terminal node, and its
usually 5-merous ovary. Adams (1962) thought that the presence of
a 5-merous ovary in H. kalmianum and in 4. H. lobocarpum indi-
cated that these species were closely related; but 5-mery in both
species appears to be derivative. The fact that H. kalmianum is
confined to once-glaciated areas (Utech & Iltis, 1970) supports the
view that it is a relatively recent derivative of H. prolificum.

3x. Hypericum kalmianum x prolificum

Utech & Iltis (1970: 335) indicate that intermediates between H.
prolificum and H. kalmianum may occur in Wisconsin, where //.
prolificum is probably always an escape from cultivation. In gardens
they seem to remain distinct; but it may be impossible to distinguish
depauperate H. prolificum plants from true H. kalmianum. (For H.
kalmianum x densiflorum see p. 102.)

4. Hypericum lobocarpum Gatt. in Bot. Gaz. 11: 275 (1886),
Tennessee FL: 29 (1887) ['labocarpum']; Sargent in Gdn Forest
10: 453, f. 57 ( 1 897); R. Keller in Engl. & Prantl, Nat. Pflanzenfam.
2nd ed. 21: 180 (1925); Small, Man. s.e.fl.: 874 (1933); Rehd.,
Man. cult, trees 2nd ed.: 639 (1940); W.P. Adams in Contr. Gray
Herb. Harv. no. 189: 12 (1962), in/ Elisha Mitchell sclent. Soc.
87: 68 (1973); Bean, Trees & shrubs hardy in Br. Isles 8th ed.: 41 1
( 1 973); Mohlenbr., ///. FL Illinois fl. pis, Hollies to Loasas: 3 1 , f. 1 2
( 1 978); Robson in Cullen et al., Eur. Gdn Fl. 4: 67 ( 1 995). Types:
U.S.A., Tennessee, Carroll Cr., Hollow Rock, August 1867 (fl &
fr), Gattinger s.n. (F*-lectotype; GH*-isolectotype); loc. cit.,
1886 (fl), Gattinger s.n. (F*, US*-syntype); 'W. Mississippi or
Tennessee', 1863, J.T. Stewart (GH-syntype).

Map 3.

? H. rostratum Raf., Fl. ludov.: 88 (1817), Herb, raf.: 55 (1833),
New Fl. 3: 95 (1836) [1838]; Eaton, Man. Bot. 6th ed.: 135
(1833). Type: unknown. This species was described by C.C.
Robin (1807) as related to H. galioides, and Rafmesque (1817)
gave a Latin translation of Robin's French description. Svenson
(1940) suggested that H. rostratum Raf. is probably an earlier
name for H. lobocarpum, a suggestion with which I agree.
Adams (1962: 50) also agreed; but he stated that, in the absence
of a type, it should be rejected (as a nomen dubium) in the
interests of nomenclatural stability. This would appear to be the
sensible course to take, and I have therefore adopted the first
undoubted name for this plant.

H. oklahomense E.J. Palmer in J. Arnold Arbor. 5: 128 (1924);
Rehd., Man. cult, trees 2nd ed.: 639 (1940). Type: U.S.A., Okla-

homa, Le Flore Co., Page, 7 October 1922 (fr), Palmer2222% (GH!-

holotype; A!-isotype).
H. densiflorum van lobocarpum (Gatt.) Svenson in Rhodora 42: 1 1

(1940); Fernald, Gray's Man. Bot. 8th ed.: 101 1 (1950); Gleason,

New Britton & Brown III. Fl. 2: 539 (1952); J.P. Gillespie in

Castanea24: 29(1958).

Icon: Sargent in Gdn Forest 10: 453, f. 57 (1897).

Map 3 Sect. 20: 4. H. lobocarpum • specimens, D records; 5. H.
densiflorum • specimens, O records; 12. H. lissophloeus A; 14. H.
chapmanii T.

Shrub 0.9-1. 5(-2) m tall, erect, with branches erect, forming large
clumps. Stems reddish, 4-lined and ancipitous when young, soon 2-
lined and rounded, becoming reddish brown and terete in 2nd
season; cortex exfoliating in strips; bark smooth, thin. Leaves sessile
or with pseudopetiole up to 5 mm long, often in axillary clusters;
lamina 35-50 x 3-10(-14) mm, narrowly oblong to oblanceolate or
linear, with margin recurved to revolute, paler or glaucous beneath,
chartaceous, deciduous at basal articulation, apex apiculate-obtuse
or apiculate-rounded to subacute, base narrowly cuneate to attenu-
ate; venation obscure beneath: c. 12-14 pairs main laterals, with
subsidiaries and densely reticulate tertiaries usually not or incom-
pletely discernible, only midrib prominent; laminar glands dense.
Inflorescence c. 5-25-flowered, without accessory flowers, with 3-
1 5-flowered dichasia from 1-3 nodes below and sometimes flowering
branches from lower nodes, the whole globose-cylindric to shortly
and broadly pyramidal; pedicels 1.5-3 mm long; bracts reduced,
elliptic to oblanceolate-spathulate or linear. Flowers 10-15 mm in
diam.; buds broadly ovoid to subglobose. Sepals 5, (3.5-)4-4.5 x
0.8-1. 5(-2) mm, subequal to equal, not enlarging but divergent to
reflexed in fruit, ± narrowly elliptic to narrowly oblong or
oblanceolate-spathulate, apiculate to acute, margins revolute, basal
veins 3-7, branching distally. Petals 5, golden yellow, becoming
incurved-deflexed, 6-7(-8) x 2.5-3.5 mm, obovate-oblanceolate
with apiculus lateral, acute. Stamens c. 100-150, longest 5-6.5 mm,
0.8-0.85 x petals. Ovary (3)4-5-merous, 2.5-3.5 x 1-1.5 mm,
narrowly ovoid, acute; placentation incompletely axile; styles (3)4-
5, 2-3 mm long, 0.8-0.85 x ovary, remaining erect, separating only
as fruit matures. Capsule 5.5-7 x 2.5-3.5 mm, narrowly ovoid-conic
to ovoid, acute to subacute, (3)4-5-lobed, exceeding sepals, thinly
coriaceous. Seeds blackish brown, 1.2-1.5 mm long, not carinate;
testa linear-reticulate. 2n = 18 (n = 9) (Hoar & Haertl, 1932).

Rocky river bottoms and banks, lake margins, swamps and open
Pinus woods; lowland up to c. 500 m.

South-eastern U.S.A. from south-eastern Oklahoma and eastern
Texas to southern Illinois, western Kentucky, eastern Alabama and
southern S. Carolina.

U.S.A. Alabama: Sumter Co., by Ala. 28, 1.2 km SE of Coatopa, 2
August 1971 (fl), Krai 43498 (BM, MO). Arkansas: Montgomery Co., P.O.
Mt Ida, 225 m, 5 July 1962 (fl), Demaree 45843 (BM); *Pulaski Co., Pulaski
Heights, Little Rock, Demaree 8207 (A, BKL, GH, NY). Illinois: Massac
Co., near Brookport, 28 July 1960, Swayne 1 104 (fide Mohlenbrock & Voigt,
1959: 242). Kentucky: *Calloway Co., between Murray and near Concord,

100

Smith & Hodgdon 4083 (GH, US); *Calloway Co., on State Line Rd. of KY
121s, 25 June 1974 (fl), Athey 2834 (MO). Louisiana: Morehouse Par., N.
side of Irvine Lake, c. 2.4 km S. of Arkansas Line NW of Beekman and c. 0.8
kmW.ofLa. 142, IVJuly 1986 (fl &fr).Da!e Thomas 974 18 (BM); Webster
Par., 6.4 kmW. of Minden, Correll & Correll 103 10 (DUKE, F, GA, MO, NY,
PH). Mississippi: *Hancock Co., 14 June 1939 (fl), Woodson & Schery 57
(MO): *Oktibbeha Co., 9.6 km W. of Starkville, Ray 8698 (MISSA). Okla-
homa: Latimer Co., SE of Damon, 1 1 August 1930 (fl), Clark 3214 (BM);
McCurtain Co., Isabel, 22 July 1915 (fl). Palmer 8082 (K). South Carolina:
Allendale Co., on 5-3-26 9.7 km S. of junction with US 301, 0.5 km W. of
junction with 5-3^3, 8 September 1967 (fl & fr), Bozeman, Radford &
Radford \ 1419 (BM, H). Tennessee: Chester Co., June 1892 (fl). Bain s.n.
(MO, Z); Corrall Co., near Hollow Rock, 1 3 August 1 897 (fr). Herb. Biltmore
3989 (BM, H, K, MO, Z). Texas: Hardin Co., 1.6-1.2 km NEof Batson, 13
November 1945 (fr), Cory 50764 (NY).

CULTIVATED. Specimens seen from the U.S.A. (1899-1920). Also
cultivated in Europe (Chittenden, 1951: 1036).

The status of 4. H. lobocarpum in relation to 5. H. densiflorum has
been debated. Gattinger originally described it as a species, but
Svenson (1940), when reducing it to varietal status under H.
densiflorum, pointed out that the two taxa could not be separated on
leaf width or habitat and that the numbers of styles and capsule valves
were only relatively different. Adams ( 1 962), on the other hand, while
agreeing that style number was not diagnostic, reported that 92. 1 % of
his H. densiflorum sample had a 3-merous ovary, whereas 79. 1 % of
his//, lobocarpum ovaries were 5-merous and a further 19.2% were 4-
merous, i.e. 98.3% were 4- or 5-merous. Taken in conjunction with
their apparently distinct distributions (separated by 240 km in Missis-
sippi/Alabama) and difference in capsule lobing, these differences
appeared to warrant recognition of these taxa as species.

Recent collections have included specimens of undoubted //.
lobocarpum from western Alabama and S. Carolina (see above),
although Adams (1973) did express doubt about the identity of the S.
Carolina population; and other specimens from Alabama tend to
have some intermediate characters (e.g. Krai 43501 (BM) from
Marengo Co., W. of Demopolis). Nevertheless the geographical
variation trends in each taxon are suggestive of original total separa-
tion and later introgression rather than incomplete separation. If this
is so, then the Alabama intermediates should be regarded as hybrids.
At any rate, it seems best to maintain these taxa as species, at least
until a field analysis of the Alabama populations is made.

4x. Hypericum lobocarpum x prolific urn

H. x dawsonianum Rehd. in Mitt. dt. dendrol. Ges. 19: 253 (1910);
W.P. Adams in Rhodora 72: 279 (1972). Type: cultivated in
U.S.A., Massachusetts, Jamaica Plain, Arnold Arboretum, 9
October 1910 (fl), Rehder s.n. (AMectotype).

Plants grown from seed of H. prolificum were found by Rehder to
have 3-5-valved lobed capsules and a multiflowered corymbose
inflorescence. Adams (1972) agreed that these plants were likely to
be hybrids with H. lobocarpum and suggested that they arose
spontaneously in the garden; and, having examined the Arnold
Arboretum specimens, I too agree with their suggested parentage.
Adams also reported field specimens intermediate between these
species from southern Missouri.

5. Hypericum densiflorum Pursh, Fl. Amer. sept. 2: 376 (1814);
Choisy, Prodr. monogr. Hyperic.: 45 (1821 ), in DC., Prodr. 1: 547
(1824); S. Watson, Bibliogr. index N. Amer. hot. 1: 125 (1878);
Coulter inBot. Gaz. 11: 84 ( 1 886), in A. Gray, Syn.fl. N. Amer. 1:
285 (1897) pro parte, excl. syn. Myriandra spathulata Spach;
Sargent in Gdn Forest 3: 525, f. 67 (1890); Svenson in Rhodora
42: 10 (1940); Rehd., Man. cult, trees 2nd ed.: 640 (1940); R.A.
Vines, Trees, shrubs & woody vines of S.W.: 756 (1960); W.P.

N.K.B. ROBSON

Adams in Contr. Gray Herb. Harv. no. 189: 14 (1962), in J.
Elisha Mitchell scient. Soc. 89: 69 (1973); Radford, Ahles &
Bell, Man. vasc.fl. Carolinas: 712(1 968); Utech & Iltis in Trans.
Wis. Acad. Sci. Arts Lett. 58: 333, f. 3 ( 1 970); R.C. Clark in Ann.
Mo. hot. Gdn 58: 209 ( 1 97 1 ); Bean, Trees & shrubs hardy in Br.
Isles 8th ed. 2: 411 (1973); Mohlenbr., ///. Fl. Illinois fl. pis,
Hollies to Loasas: 37, f. 16 (1978); Godfrey & Wooten, Aquatic
& wetland pis ofs.e. U.S., Dicots: 346, f. 157 (1981); N. Robson
in Cullen et al., Eur. Gdn Fl. 4: 67, ff. 10.1, 10.1 1 (1995). Type:
U.S.A., Virginia, on the dry ridges and savannahs of the Virginia
mountains, Pursh s.n. (OXF?-holotype) - see Svenson (1940).
Map 3.

H. prolificum sensu Torrey & Gray, Fl. N. Amer. 1: 159 (1838) pro
parte quoad syn. H. densiflorum Pursh.

H. rosmarinifolium sensu Torrey & Gray, Fl. N. Amer. 1: 1 59 ( 1 838);
R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 180
( 1 925); non Lam. (1797).

H. prolificum var. densiflorum (Pursh) A. Gray, Manual 2nd ed.: 50
(1856).

H. glomeratum Small in Bull. N. Y. hot. Gdn 1: 281 (1899) pro parte
quoad typum, Man. s.e. fl.: 87 (1933). Type: U.S.A., North
Carolina, Watauga Co., Grandfather Mtn, August 1 896 (fr), Huger
s.n. (NY-holotype).

H. interior Small in Bull. Torrey hot. Club 28: 359 (1901). Type:
U.S.A., Tennessee, Jefferson Co., prope Dandridge, July 1842
(fl), Rugel 154 (NY-holotype; BM!, G!, K!, W!-isotypes).

H. nothum Rehd. in Mitt. dt. dendrol. Ges. 19: 254 ( 1 9 1 0) qua hybr.,
Man. cult, trees 2nd ed.: 639 (1940); W.P. Adams in Rhodora 74:
280 (1972); Bean, Trees & shrubs hardy in Br. Isles 8th ed. 2: 416
(1973). Type: cultivated in U.S.A., Massachusetts, Jamaica Plain,
Arnold Arboretum, 10 October 1910 (fr), Rehder s.n. (A!-
lectotype, selected here). Rehder interpreted his specimens as
hybrid (H. densiflorum x kalmianum) on account of the occur-
rence of a few 4- or 5-styled capsules among the predominantly
3-styled ones. Despite these, however, as Adams pointed out, the
specimens can readily be assigned to H. densiflorum.

H. revolutum R. Keller in Bot. Jb. 58: 194 (1923), in Engl. & Prantl,
Nat. Pflanzenfam. 2nded. 21: 180(1925); non Vahl (1790). Type:
U.S.A., Georgia, Whitfield Co., Spring Creek near Sarnell, 225
m, 21 December 1903 (fr), Harper 2032 (Bf-holotype; F!, GH!,
NY-isotypes).

Icones: Sargent in Gdn Forest 3: 527, f. 67 ( 1 890); Mohlenbr., ///. Fl.
Illinois fl. pis, Hollies to Loasas: 39, f. 16 (1978).

STUDIES IN HYPERICUM

101

ShrubQ.6-3 m tall, erect, with branches ± erect, rather stiff, numerous,
forming rather slender bush, but with adventitous shoots from roots
sometimes producing ± extensive thickets. Stems green to reddish, 4-
lined and ancipitous when young, soon 2-lined and rounded, becoming
reddish brown and terete in 2nd season; cortex exfoliating in strips;
bark smooth, thin. Leaves sessile, sometimes in immature clusters in
leaf axils; lamina 20^5 x 2-7 mm, very narrowly elliptic-oblong or
oblanceolate to linear, with margin recurved to revolute, paler and
often glaucous beneath, chartaceous, deciduous at basal articulation,
apex apiculate or apiculate-obtuse to subacute, base narrowly cuneate
to attenuate; venation sometimes obscure beneath: c. 14-17 pairs
main laterals, with subsidiaries and tertiary reticulation sometimes
visible, only midrib prominent; laminar glands dense. Inflorescence c.
5-25-flowered, without accessory flowers, with (2-)5-15-flowered
dichasia from 1-2 nodes below and sometimes flowering branches
from lower nodes, the whole broadly pyramidal to broadly cy lindric or
obpyramidal; pedicels 0.5-5 mm long; bracts reduced, oblanceolate-
spathulate to linear. Flowers 10-17(-20?) mm in diam.; buds broadly
ovoid-ellipsoid. Sepals 5, (4-)4.5-6x 1-1. 5 mm, unequal or subequal,
not enlarging but divergent to reflexed in fruit, narrowly oblong-
lanceolate or narrowly oblong to oblanceolate-spathulate, subacute or
apiculate to acute, margins revolute, basal veins 1-3, the laterals
sometimes branched. Petals 5, deep golden yellow, becoming
(incurved-) deflexed, 6-9 x 2.5-3.5 mm, obovate-oblanceolate with
apiculus lateral, acute. Stamens c. 100-150, longest 4. 5-7 mm, c. 0.8
x petals. Ovary 3^(-5)-merous, 3-3.5 x 1-1.3 mm, narrowly
pyramidal-ovoid, acute; placentation incompletely axile; styles 3-
4(5), 2-3 mm long, 0.7-0.85 x ovary, remaining erect, separating only
as fruit matures. Capsule 5-6(-7) x 2-3 mm, narrowly ovoid-conic or
narrowly ovoid to cylindric-ovoid, acute, not or scarcely lobed,
exceeding sepals, thinly coriaceous. Seeds reddish brown, 0.8-1.3
mm long, not carinate; testa linear-reticulate. 2n = 1 8 (n = 9) (Adams
in Robson & Adams, 1968). Adams also reported 2n = 27 (n = 1 1-14)
in one population due to unequal segregation (Adams, 1959)

Wet or moist habitats (meadows, lake margins, open stream banks,
pinelands, bogs, ditches), also on dry road embankments and rocky
hillsides; lowland to c. 1000 m.

Eastern U.S.A. from central Georgia northward along the coastal
plain to New Jersey and thence southwestward along the Appala-
chian Mts from Pennsylvania to northern Georgia and central
Alabama.

U.S.A. Alabama: *Jefferson Co., Shade's Mt. near Birmingham, Leeds
2140 (PH); Tuscaloosa Co. (see Adams, 1962). Delaware: Sussex Co.,
Seaford, Nauticoke R., 31 August 1882 (fr), Commons s.n. (K). Georgia:
Bartow Co., Rydal, N. of Cartersville, 1 1 July 1958 (fl), Adams 77 (K, MO);
*Catoosa Co., east-flowing tributary to Chickamauga Creek, 5 km NE of
Ringgold, 240 m, 26 June 1948 (fl), Cronquist 5390 (MO). Maryland:
Wiscomico Co., Salisbury, June pre-1890 (fl & fr), Canby s.n. (K). New
Jersey: Burlington Co., 3.2 km W. of Warren Grove, 18 June 1948 (fl),
Lawrence & Dress 264 (BM); Ocean Co., Bayville, 3 1 August 1937 (fl & fr),
Moldenke 10163 (BM). New York: ? Co., Long Island, fide Adams (1959:
map 6). North Carolina: Avery Co., 6.2 km W. of Allegany Co. line on NC 88,
30 July 1 979 (fl & fr), Leonard & Russ 2644 (BM, H); Macon Co., Highlands,
6th Street, 31 July 1975 (fl), Bouffon & Wood 1775 (H, MO). Pennsylvania:
Fagette Co., Ohiopyle, 15 October 1933, Jennings s.n. (CA). South Carolina:
Georgetown Co., 8 km W. of Georgetown, 3 August 1939 (fl), Godfrey in PI.
Exs. of Gray 1371 (BM, K). Tennessee: Knox Co., Big Ridge State Park, 4
July 1954 (fl), Norm 18622 (BM); Rhea Co., near Watts Bar Dam, 25 June
1947 (fl), Shanks, Sharp & Clebsch 4197 (BM). Virginia: ? Co., Tygart's
valley river, 1843 (fl & fr), Gray & Sullivant s.n. (K); Grayson Co., Blue
Ridge Parkway, 0.5 km N. of Route 89, 19 July 1958 (fl), Adams 109 (K,
NCU*). West Virginia: Randolph Co., below Elkins, 24 August 1907 (fl),
Render s.n. (K, MO); *Webster Co., Camp Caesar near Cowen, 1 8 June 1 936
(fl), Williams 490 (BKL, DUKE, F, MO, PH, SMU, TENN, US, WVA).

Despite the existence of some intermediate populations, it seems
best to maintain 5. H. denslflorum as specifically distinct from 4.
H. lobocarpum (q.v.). H. densiflorum usually has a trimerous
ovary, but 4- or 5-merous ovaries may also be present on the same
plant. The capsule is usually but not always unlobed. However,
except for members of the intermediate Alabama populations, it is
nearly always possible to allocate a specimen to one or other
species.

Whereas the southwesternmost (Alabama) populations of H.
densiflorum verge morphologically towards H. lobocarpum, some in
Tennessee and northern Georgia with narrow leaves and narrow,
small-flowered inflorescences (H. interior Small) tend towards H.
galioides. H. densiflorum is apparently always distinguishable in the
wild from 2.H. prolificum by the smaller, more numerous flowers amd
fruits in a shorter and relatively broader inflorescence. In cultivation,
however, these distinctions may be far less clear (see below).

Adams (1973) reported evidence of poor seed production in some
populations, a condition that, he suggested, might be correlated with
the occurrence of triploidy (see above).

5x. Hypericum densiflorum x galioides?

H. x arnoldianum Rehd. in Mitt. dt. dendrol. Ges. 19: 253 (1910);
W.P. Adams in Rhodora 74: 276 (1972). Type: cultivated in
U.S.A., Massachusetts, Jamaica Plain, Arnold Arboretum, 9 Oc-
tober 1910 (fr), Rehder (AMectotype, selected here).

Render's belief that the parentage of this hybrid was H. galioides x
lobocarpum was not substantiated by Adams (1972), who found only
a few well-formed seeds in each fruit. He argued that, whilst the
partially lobed fruit and the 1 6 per cent of fruits with 4 styles indicated
that H. lobocarpum was one parent, there were no characters of H.
galioides present. On the other hand, characters of the Tennessee
populations of//, densiflorum were present; and so he suggested that
the most likely parentage of H. x arnoldianum was H. densiflorum
x lobocarpum. Support for this hypothesis had been provided earlier
by Hoar & Haertl (1932), who reported 'no irregularity in chromo-
somebehaviour nor morphological sterility of pollen' when recording
the chromosome number n = 9, and concluded that this hybrid
'apparently came from compatible parents' (which //. densiflorum
and H. lobocarpum almost certainly would be).

On the other hand, Rehder's original plants came 'from seed of//.
galioides' (specimen 9x191 0). Adams suggested that Rehder might
have misidentified the relevant specimen(s) of H. galioides; and
since the above-mentioned Tennessee population of H. densiflorum
is morphologically the most similar part of that species to //.
galioides, he may well be right. After an examination of Rehder's
specimens, however, I cannot be sure that H. galioides is not
involved; and specimens grown subsequently at the Arnold Arbore-
tum from plants obtained from Cole Nursery, Painesville, Ohio (AA
856-38 (MO)) look like //. densiflorum (or possibly H. lobocarpum)
except for a narrower, more elongated inflorescence (from 3 nodes)
that is reminiscent of that of H. galioides.

5xx. Hypericum densiflorum x prolificum

In gardens, at least in the British Isles, it is not always easy to decide
whether a plant belongs to one or the other of the above two species.
If the specimen has small flowers and relatively narrow leaves but
the inflorescence of H. prolificum, then it can probably be regarded
as an impoverished example of that species (cf. H. foliosum sensu
Jacquin, PL hort. schoenbr. 3: t. 299, 1798). If it has a shorter,
broader inflorescence and smaller leaves than those of//, prolificum,
and hybridization with H. frondosum or H. kalmianum can be ruled

102

N.K.B. ROBSON

out, then it could quite possibly be the hybrid H. densiflorum x
prolificum. A low (c. 0.4-0.6 m tall) compact shrub grown in British
gardens as H. x arnoldianum could belong here but is more likely to
be//, densiflorum xlobocarpum or even//, densiflorum xkalmianum
(the alleged parentage of H. nothum Render).

6. Hypericum galioides Lam., Encycl. 4: 16(1797); W\\ld., Sp. pi.
3: 1451 (1802); Pursh, Fl. Amer. sept. 2: 376 (1814); Elliott,
Sketch hot. S. Carolina 2: 28 (1821); Choisy, Prodr. monogr.
Hyperic. : 52 ( 1 82 1 ), in DC., Prodr. 1 : 550 ( 1 824); Torrey & Gray,
Fl. N. Amer. 1: 159 (1838); S. Watson, Bibliogr. index N. Amer.
hot. I: 126 (1878); Coulter in flo/. Gaz. 11: 85 (1886), in A. Gray,
Syn.fl. N. Amer. 1: 286 (1897); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nded.21: 181 ( 1925); Svenson in/?/zoJ0ra 42: 12,
t. 587 f. 3 (1940) pro parte, quoad van typicum; Rehder, Man.
cult, trees & shrubs 2nd ed.: 639 (1940); Gillespie in Castanea
24: 29 (1958); R.A. Vines, Trees, shrubs & woody vines ofS.W.:
757 (1960); W.P.Adams inContr. Gray Herb. Harv. no. 189: 171
( 1 962), in J. Elisha Mitchell scient. Soc. 89: 69 ( 1 973); Correll &
Johnston, Man. vase. pis. Texas: 1063 (1970); R.C. Clark in Ann.
Mo. hot. Gdn 58: 209 (1971); R. Long & Lakela, Fl Trap.
Florida: 607 (1971); Bean, Trees & shrubs hardy in Br. Isles 8th
ed. 2: 414 (1973); Godfrey & Wooten, Aquatic & wetland pis s.e.
U.S., Dicots: 346, f. 1 56 ( 1 98 1 ); Clewell, Guide vase, pis Florida
Panhandle: 372 (1985); Godfrey, Trees, shrubs & woody vines n.
Florida, etc.: 368, f. 177 (1988); N. Robson in Eur. Gdn Fl. 4: 69
(1995). Type: U.S.A., S. Carolina ('Carol, merid.'), no precise
locality, Fraser (P-LA-holotype, BM!-microfiche; G-DC!-
isotype; GH-sketch). Svenson ( 1 940: 1 3) stated that this collection
is apparently identical with several from near Wilmington, N.
Carolina, although 'merid.' implies S. Carolina.

Fig. 15 A, Map 4.

H. axillare Lam., Encycl. 4: 161 (1797). Type: U.S.A., no precise
locality, ? 'in Herb. D. de Jussieu' (P-holotype, BM! -microfiche;
GH*-isotype).The specimen labelled as//, axillare in the micro-
fiche series of the de Jussieu herbarium appears to be//, prolificum.
The GH isotype is a fragment (fide Adams, 1962).
H. triplinerve Vent., Desc. pi. nouv.: 58 & t. ( 1 800); Pursh, Fl. Amer.
sept.: 378 (1814). Type: U.S.A., cult, ex 'sur lesbordsdel'Ohio',
n.d. (fl & fr), Michaux s.n. (G-holotype). The locality cited by
Ventenat is well outside the present distribution of H. galioides
and must surely be an error.

H.fasciculatumMichyi.exWi\ld., Sp.pl. 3: 1452(1 802); Michx.,F/.
bor.-amer. 2: 80 (1 803); non Lam. (1797). Type: U.S. A., Carolina,
Michauxl s.n. (P-holotype, BM! -microfiche).
H. michauxii Poir., Encycl., Suppl. 3: 696 (1813). Type as for H.
fasciculatum Michx. ex Willd., non Lam.

H. fasciculatum [van] (3 sensu Choisy, Prodr. monogr. Hyperic.: 59
(1821) pro parte, quoad syn.

H. ambiguum Elliott, Sketch hot. S. Carolina 2: 30 ( 1 82 1 ); Torrey &
Gray, Fl. N. Amer. 1: 62 (1838); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 181 (1925); Small, Man. s.e. fl.: 871
(1933). Type: U.S.A., S. Carolina, near Columbia, Elliott s.n.
(CHARL-holotype).

H. rosmarinifoliumt sensu Elliott, Sketch hot. S. Carolina 2: 29
(1821), non H. rosmarinifolium Lam. (1797).

Myriandra galioides (Lam.) Spach, Hist. nat. veg. Phan. 5: 437
(1836), in AnnlsSci. nat. (Bot.) II, 5: 365 (1836); K. Koch, Hort.
dendroi.: 66 (\853).

M. michauxii (Poir.) Spach, Hist. nat. veg. Phan. 5: 437 (1836), in
Annls Sci. nat. (Bot.) II, 5: 365 (1836).

M. prolifica [var]. P ramosa K. Koch, Hort. dendroi: 66 (1853).
Type as for '//. galioides Pursh', i.e. H. galioides Lam.

Brathydium ambiguum (Elliott) K. Koch, Hort. dendroi.: 66 (1853).

Hypericum galioides var. ambiguum (Elliott) Chapm., Fl. South. U.
S.: 40 (1865); S. Watson, Bibliogr. index N. Amer. hot. 1: 127
( 1 878). Type: Chapman cites '//. ambiguum Ell.? Torrey & Gray',
but Torrey & Gray ( 1 838) do not mention var. ambiguum, Chap-
man's name must therefore have Elliott's type.

H. galioides var. axillare (Lam.) Griseb., Cat. pi. Cub. : 39 ( 1 866) pro
parte, quoad typum.

H. galioides [var.] pallidum Mohr in Contr. U. S. natn. Herb. 6: 621
(1901); Lott in / Arnold Arbor. 19: 149 (1938); Svenson in
Rhodora 42: 14, t. 587 f. 4 (1940), nom. illegit. (Art. 63). Type as
for '//. galioides var. ambiguum Chapman nonambiguum Elliott' ;
but the type nevertheless is that of Elliott, see above.

H. spathulatum R. Keller in Bot. Jb. 58: 195 (1925), in Engl. &
Prantl, Nat. Pflanzenfam. 2nd ed. 2: 180 (1925), non (Spach)
Steud. ( 1 840). Type: U.S.A., Georgia, Lee Co., Flint R., Muckalee
Swamp, 1 1 July 1901 (fr), Harper 1 155 (Bf-holotype; A!, BM!,
GH*, NY*, US!-isotypes).

Icones: Bean in Gdnrs Chron. Ill, 24: 301, f. 88 (1898); Godfrey &
Wooten, Aquatic & wetland pis s.e. U. S., Dicots: 347, f. 1 56 ( 1 98 1 ).

Map 4 Sect. 20: 6. H. galioides
specimens, D records.

specimens, O records; 8. H. lloydii

Shrub 0.5-1.5 m tall, erect, with branches strict to ascending,
forming rounded clumps. Stems stramineous to reddish, 6-lined and
ancipitous when young, soon 4-lined and rounded, then terete;
cortex exfoliating in small roughish flakes to reveal fine lattice of
laticifers; bark smooth, thin. Leaves sessile or apparently
pseudopetiolate, with those in axillary immature clusters sometimes
almost as large; lamina 15-32(-37) x 1-7 mm, very narrowly
oblong-elliptic or oblanceolate to linear, with margin recurved to
revolute, paler but not glaucous beneath with lamina visible on both

STUDIES IN HYPER/CUM

103

Fig. 15 A. H. galioides: (a) habit; (b) leaf (upper half); (c) sepal; (d) petal; (e) stamen; (f) ovary; (g) fruit. B. H. tenuifolium: (h) habit. C. H. lloydii: (i)
habit (a, h, i x !/2; b x 3: c-g x 4). A. Rugel 495. B. Herb. Biltmore 2563a. C. Eggert s.n.

104

N.K.B. ROBSON

sides of midrib, chartaceous, deciduous at basal articulation, apex
rounded or apiculate-obtuse to acute, base attenuate; venation (some-
times obscure): numerous main laterals and subsidiaries, with tertiary
reticulation often visible, only midrib prominent; laminar glands
dense; inframarginal glands dense. Inflorescence 3-c\ 15-flowered,
without accessory flowers, with ( 1 )3-5-flowered dichasia from 3^
nodes below and often flowering branches from lower nodes, the
whole narrowly cylindric; pedicels 0.5-1 mm long; bracts somewhat
reduced, foliar. Flowers 9-14 mm in diam.; buds narrowly ovoid.
Sepals 5, 3.5-6.5 x 0.5-1 .5 mm, subequal to equal, not enlarging or
spreading in fruit, oblanceolate-spathulate to linear, acute to apiculate-
obtuse, 1 -veined, deciduous. Petals 5, bright yellow, becoming
somewhat deflexed, 5-9 x 2-5.5 mm, obovate-oblanceolate with
apiculus lateral, acute. Stamens c. 60-120, longest 3.5-7 mm, c.
0.65 x petals. Ovary 3-merous, 2-3 x 0.5-1 mm, very narrowly
pyramidal-ovoid, acute; placentation parietal; styles 3, 4-5 mm
long, 1.2-1.35 x ovary, remaining erect, separating only as fruit
matures. Capsule 4.5-6 x 2.5-3.5 mm, ± narrowly ovoid-conic,
shorter than sepals, thinly coriaceous. Seeds dark brown, 0.7-0.8
mm long, carinate?; testa finely reticulate. 2n = 1 8 (n = 9) (Adams in
Robson & Adams, 1968).

Wet or moist, open habitats (stream banks, swamps, river bottoms,
flood plains, lake margins, roadside ditches, low pine forest); coastal
plain, below 200 m.

South-eastern U.S.A. from North Carolina to northern Florida and
west to eastern Texas, excluding most of the Mississippi delta.

U.S.A. Alabama: *Covington Co., c. 1 1 .2 km SSE of Carolina, by co.e
31,3 October 197 1 (fr). Krai 44760 (MO); Houston Co., Indigo Pond SE of
Cottonwood, 1 August 197 1 (fr), Krai 43399 (BM). Arkansas: Union Co., E.
of Strong, 7.5 km W. of Ouachita R., Felsenthal Nat. Wildlife Refuge, 22
October 1987 (fr), Dale Thomas 103014 (MO). Florida: Columbia Co., Lake
City, 1 1-19 July 1895 (fl), Nash 2190 (K); Gulf Co., shores of Dead Lake
near Wewahitchka, 3 September 1958 (fr), Godfrey 57585 (BM, FSU*).
Georgia: Lowndes Co., E. side of Little River, c. 8 km W. of Hahira, 19 June
1958 (fl), Adams 50 (K); Dooly Co., bank of Flint R., 57 m, 1 1 July 1901 (fl),
Harper 1061 (BM). Louisiana: *Calcasieu Par., L. Charles, 10 September
1 898 (fr), McKenzie 5 1 7 (MO); New Orleans, 1 832 (fr), Drummond 50 (BM,
K); Natchitoches Par., 25 May 1941 (fl), Keefe s.n. (H). Mississippi: Perry
Co., outskirts of Beaumont, 16 July 1950, Webster & Wilbur 3409 (G); * ?
Co., lower Pearl River, May 1859 (fl), Hilgard s.n. (MO). North Carolina:
Columbus Co., Co. Rt. 1928 SE of Old Dock near Juniper Creek, 1 1 July
1968 (fl), Leonard 1754 (BM, H, Z); Pender Co., N. of Burgaw, near U.S.
1 17, 23 July 1966 (fl), Bradley 3333 (BM, H). South Carolina: Colleton Co.,
8km W. of junction of SC64 and SC641, 14 July 1958 (fl), A dams 89 (K);
*Williamsburg Co., 8 km S. of Kingstree, 10 July 1959 (fl), Godfrey & Tryon
383 (MO). Texas: Hardin Co., 16 km SE of Votaw, 23 July 1978 (fl), Fryxell
3015 (BM); *Harris Co., Sheldon, 1903 (fl & fr), Reverchon 3828 (MO).

The leaves of H. galioides vary in width from that of typical H.
densiflorum ('var. ambiguum') to as narrow as those of H. nitidum;
but the lamina (except for the midrib) is always thin and visible
beneath, at least in the living plant. The long narrow inflorescence
distinguishes it from H. densiflorum but not from H. nitidum, which
is a larger plant, having leaves usually with no lamina visible
beneath.

As Adams (1962) has pointed out, the names H. galioides Lam.
and H. axillare Lam. were published in the same work. The early
nineteenth century botanists were unsure of the application of H.
axillare, so H. galioides became the established name.

7. Hypericum tenuifolium Pursh, Fl, Amer. sept.: 377 (1814).

Type: U.S.A., Georgia, no precise locality (fl), Enslen s.n. (K-

holotype?; PH-isotype).
Figs 15B, 16E, Map 5.

Map 5 Sect. 20: 7. H. tenuifolium • specimens, O records (incomplete,
as Adams (1959) did not differentiate Spp. 7 and 1 1); 15. H. buckleyi •
specimens, D records.

H. com? sensu Walter, Fl. Carol.: 190 (1788), non H. coris L.
(1753). Type': U.S.A., no precise locality, Herb. Walter (BM\).

H. fasciculatum [var.] P sensu Choisy, Prodr. monogr. Hyperic.:59
( 1 82 1 ) pro parte, quoad syn. H. tenuifolium; Torrey & Gray, Fl. N.
Amer. 1: 160 (1838) pro parte, excl. syn. H. axillare.

H. fasciculatum var. laxifolium Choisy in DC., Prodr. 1: 554 (1824)
pro parte, excl. syn. H. michauxii.

H. fasciculatum var. aspalathoides sensu Torrey & Gray, Fl. N.
Amer. 1: 672 (1840) pro parte, excl. syn.Myriandrabrachyphylla,
non H. aspalathoides Willd. (1802), nom. illegit. (Art. 63).

H. aspalathoides sensu Small, Man. s.e. fl.: 872 (1933) pro parte;
R.A. Vines, Trees, shrubs & woody vines ofS.W.: 759 (1960).

H. galioides var. reductum Svenson in Rhodora 42: 14 (1940) pro
parte excl. typum. Type as for H. aspalathoides Willd.

H. reductum W.P. Adams in Contr. Gray Herb. Harv., no. 189: 31
(1962), in J. Elisha Mitchell sclent. Soc. 89: 70 (1973); Radford,
Ahles & Bell, Man. vasc.fl. Carolinas: 712 (1968); R.C. Clark in
Ann. Mo. hot. Gdn 58: 209 (1971); R. Long & Lakela, Fl. Trop.
Florida: 608 (1971); Godfrey & Wooten, Aquatic & wetland pis
s.e. U.S., Dicots: 343, f. 155 (1981); Clewell, Guide vase, pis
Florida Panhandle: 372 (1985); Godfrey, Trees, shrubs & woody
vines n. Florida, etc.: 358, f. 171 (1988). Type: U.S.A., N.
Carolina, Wilmington, n.d. (fl), Curtis s.n. (GH!-holotype, NY-
isotype).

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 358, f.
171 (1988).

STUDIES IN HYPERICUM

105

Shrub 0. 1-0.5 m tall, decumbent, with branches interweaving at the
base but not rooting, forming low mats, or sometimes more erect in
dense vegetation. Stems reddish, 6-lined when young, eventually 4-
lined, then terete; cortex exfoliating in strips or flakes; bark smooth,
thin. Leaves sessile, 4-1 1 x 0.4-0.8(-1?) mm, with those in axillary
clusters equal or shorter, linear-subulate, dull green above, with
margin revolute, completely concealing all but midrib beneath,
chartaceous, deciduous at articulation below prominent midrib base
(cf. 11. brachyphyllum), apex rounded with prominent hydathode,
base parallel or slightly expanded; midrib unbranched; glands dense,
in 2 rows, visible beneath only. Inflorescence 1-7-flowered, without
accessory flowers, with l(3)-flowered dichasia from up to 4 nodes
below, rarely with one pair of flowering branches, the whole ±
narrowly cylindric; pedicels absent or very short; bracts foliar.
Flowers 10-14 mm in diam.; buds narrowly cylindric-ellipsoid.
Sepals 5, (2-)2.5— 4 x 0.4-0.5 mm, unequal, linear-subulate, acute to
rounded, 1 -veined. Petals 5, bright? yellow, spreading, 5-10 x 2-5
mm long, oblanceolate-oblong to obovate, with apiculus lateral,
acute. Stamens c. 50-90, longest 4.5-8 mm, c. 0.85-0.9 x petals.
Ovary 3-merous, 2-3 x 0.5-0.8 mm, very narrowly cylindric, acute;
placentation parietal?; styles 3, 2-3 mm long, 0.9-1.2 x ovary,
usually separating in fruit. Capsule (4-)5. 7-9.5 x 1 .5-2 mm, nar-
rowly (sub)cylindric, exceeding sepals, thinly coriaceous. Seeds
blackish, c. 0.5 mm long; testa coarsely reticulate (alveoli square to
hexagonal). 2n = 18 (n = 9) (Adams in Robson & Adams, 1968).

Dry sandy woods, dunes and dune hollows; lowland and coastal.

South-eastern U.S.A. from North Carolina to central Florida and
southern Alabama; four distinct populations: 1) south-eastern N.
Carolina and adjacent S. Carolina; 2) extreme south-eastern S.
Carolina and adjacent eastern Georgia; 3) central peninsular Florida;
4) coastal Florida 'panhandle' and adjacent Alabama.

U.S.A. Alabama: Baldwin Co., by Ala 180 towards Fort Morgan, c. 24
km W. of junction with Ala 59, 8 June 1971 (f\),Kral 43098 (BM, MO); Pike
Co., Spring Hill, 5 August 1897 (fr), Bush 348 (MO). Florida: Bay Co., Gulf
Lagoon Beach, SW of Panama City, 20 May 1961 (e. fl), Adams 776 (K);
Franklin Co., Dog Island, 17 June 1970 (fl), Godfrey 69546 (H); Lake Co.,
Ocala National Forest, 1.75 km N. of Wildcat Lake, c. 3.2 km E. of Marion/
Lake county line, 2 August 1962 (fl), Ward & Will 3048 (BM); Lee Co., near
Coconut, 14 April 1930 (fl), Moldenke 965 (K); St. Lucie Co., 10. 1 km S. of
White City, 1 2 April 1950 (fl), Hansen924 (BM). Georgia: Bryan Co., E. side
of Canochee R.c. 13 km E. of Pembroke, 8 October 1960 (fr), Adams! 17 (K,
MO); *McIntosh Co., 3.2 km W. of Eulonia, 9 October 1960 (fr), Adams 723
(MO). North Carolina: Brunswick Co., SE of Wilmington, 25 June 1 890 (fl),
Cavilled (K); New Hanover Co., Carolina Beach, 1 8 June 1939 (fl), Godfrey
1260 (BM, K). South Carolina: * ? Co., Bluffton, 1 874 (fl), Mellenchamp s.n.
(MO).

H. tenuifolium, like H. lloydii, is closely related to (probably derived
from) H. galioides, with which it shares all but the peninsular
Florida part of its area. It is quite distinct from H. galioides, however,
in habit, leaf size, inflorescence and habitat. Its distribution does not
overlap the piedmont area of H. lloydii, which is also decumbent in
form but has rooting stems and shorter leaves and (usually) sepals.
For a comparison between H. tenuifolium and H. brachyphyllum, see
p. 110.

Adams (1962) was aware that H. tenuifolium Pursh might be the
earliest name for this species. Having been unable to find an
authentic specimen, however, he published a new name, H. reductum.
This was no doubt intended to maintain Svenson's (1940) epithet,
although, because Svenson cited H. aspalathoides Willd. in syn-
onymy, Adams could not treat his name as a stat. et comb. nov. In
Kew (K), however, there is an Enslen specimen from Herb. Pursh.
which I consider to be an authentic specimen of H. tenuifolium, and
so the Philadelphia (PH) specimen cited by Adams can also be

considered as authentic. Both these specimens belong to Adams's H.
reductum, a name which must unfortunately therefore pass into
synonymy under H. tenuifolium Pursh.

8. Hypericum lloydii (Svenson) W.P. Adams in Contr. Gray Herb.
Harv. no. 189: 32 (1962), in 7. Elisha Mitchell scient. Soc. 89: 70
(1973); Radford, Ahles & Bell, Man. vase. fl. Carolinas: 712
(1968); R.C. Clark in Ann. Mo. hot. Gdn 58: 209 (1971). Type:
U.S.A., S. Carolina, Aiken Co., Graniteville, 4 August 1 898 (fl),
Eggert s.n. (NY-holotype; F!, MO, US-isotypes).

Fig. 15C, Map 4.

H. galioides var. lloydii Svenson [in Rhodora 42: t. 587 f. 8 (1940)
nomen] in Rhodora 54: 207 (1952).

Icon: Svenson in Rhodora 42: t. 587 f. 8 (1940).

Shrub 0.1-0.5 m tall or rarely taller, with branches straggling and
rooting, forming low rounded clumps or mats. Stems reddish, 6-
lined and ancipitous when young, sometimes eventually 4-lined,
then terete; cortex exfoliating in strips or flakes; bark smooth, thin.
Leaves sessile, with those in axillary clusters smaller; lamina 13-25
x 0.5-0.8 mm, linear-subulate, with margin revolute, paler but not
glaucous beneath, with lamina sometimes visible on both sides of
midrib, chartaceous, deciduous at basal articulation, apex rounded
to retuse with prominent hydathode, base parallel; midrib unbranched;
laminar glands rather dense, visible beneath only. Inflorescence 1-
3-flowered, without accessory flowers, with l-3(?-5)-flowered
dichasia from up to 5 nodes below, without flowering branches, the
whole narrowly pyramidal; pedicels c. 0.5 mm long; bracts foliar.
Flowers c. 12-14 mm in diam.; buds broadly ovoid. Sepals 5, (3-)
4.5-7 x 0.5-0.8 mm, unequal, linear-subulate, subacute to rounded,
1 -veined. Petals 5, golden yellow, spreading, 5-7.5 x 3^4 mm,
oblanceolate-oblong with apiculus obtuse. Stamens c. 100, longest
(5-)6-7 mm, almost equalling petals. Ovary 3-merous, c. 3 x 1 mm,
narrowly triangular-ovoid; styles 3, 2.5-3 mm long, 0.8-1 x ovary,
separating in fruit or earlier; placentation parietal? Capsule 3-4 x 2-
2.5 mm, 'ovoid', shorter than sepals, thinly coriaceous. Seeds black,
carinate?, 0.7 mm long; testa not seen.

Dry woods, pine forest, granite outcrops, roadside embankments;
inner coastal plain and piedmont; c. 150-300 m.

South-eastern U.S.A. from North Carolina to Alabama, excluding
Florida.

U.S.A. Alabama: *Tallapoosa Co., Harper 3691 (GH. PH. US). Geor-
gia: *Richmond Co., August, 17 July 1 899, Cuthbert s.n. (FLAS, NY). North
Carolina: Chatham Co., U.S. 15-501, c. 13km S.of Pittsboro, 30 June 1966
(fl), Bell 18577 (H); *Granville Co., Oxford, Gillespie 394 (DUKE, FSU.
NCSC); *Wake Co., Gary, July 1898 (fl), Ashe s.n. (MO). South Carolina:

106

N.K.B. ROBSON

Map 6 Sect. 20: 9. H. nitidum: a. subsp. cubense • specimens, O
records; b. subsp. nitidum • specimens, D records; c. subsp. exile A ;
10. H. limosum V.

Lancaster Co., 3. 2 km S.ofTaxahaw,40Acre Rock, 19April 1 964 (st), Creem
383 (BM).

H. lloydii is closely related to 6. H. galioides but differs from it in
habit, leaf-shape and habitat (both in altitude and in favouring drier
sites); and these species occupy almost distinct areas. It therefore
merits recognition as a species.

9. Hypericum nitidum Lam., Encycl. 4: 1 60 ( 1 797); Choisy, Prodr.
monogr. Hyperic.: 59 (1821), in DC., Prodr. 1: 554 (1824); W.P.
Adams in Contr. Gray Herb. Harv. no. 189: 26 (1962), in /
Elisha Mitchell sclent. Soc. 89: 69 (1973); Radford, Ahles &
Bell, Man. vasc.fl. Carolina*: 1 1 (1968); R.C. Clark in Ann. Mo.
hot. Gdn 58: 209 ( 1 97 1 ); Godfrey & Wooten, Aquatic & wetland
pis s.e. U.S. Dicots: 345 ( 1 98 1 ); Clewell, Guide vase, pis Florida
Panhandle: 372 (1985); Godfrey, Trees, shrubs & woody vines n.
Florida, etc.: 363, f. 174 (1988). Type: U.S.A., Carolina, ? (P-
holotype, microfiche!, GH-fragm. & photograph).

Map 6.

Myriandra nitida (Lam.) Spach, Hist. nat. veg. Phan. 5: 435

( 1 836), in Annls Sci. nat. (Bot.) II, 5: 365 ( 1 836); K. Koch, Hon.

dendrol.: 66 (1853).
Hypericum fasciculatum sensu Torrey & Gray, Fl. N. Amer. 1: 672

(1840); Trevir., Hyper, sp. animadv.: 15 (1861); Coulter in Bot.

Gaz. 1 1: 85 ( 1 886); R. Keller in Engler & Prantl, Nat. Pflanzenfam.

2nd ed. 21 : 1 80 ( 1 925); et auct. plur. pro parte omnes, quoad syn.

H. nitidum Lam.
H. galioides \ar.fasciculatum (Lam.) Svenson in Rhodora 42: 12, t.

587, ff. 1-2 (1940) pro parte excl. typum.

Shrub or small tree (0.3-)0.6-4.5 m tall, erect, with branches erect to
ascending, forming thickets. Stems reddish, 4-lined and ancipitous
when young, becoming narrowly 2-winged, eventually terete; cortex
exfoliating in flakes or narrow strips; bark brown or reddish to grey,
smooth, thin. Leaves sessile, (6-) 10-21 (-26) x 0.5-1.4 mm, with
those in axillary clusters as long or usually somewhat shorter, linear

or linear-subulate, sometimes slightly broadened distally, with mar-
gin revolute and often completely concealing all but the (sometimes
slightly raised) midrib beneath, the non-midrib area sometimes
papillose, chartaceous to subcoriaceous, deciduous at basal articula-
tion, apex rounded with prominent hydathode to obtuse-apiculate or
long acuminate, base parallel or narrowly cuneate; midrib
unbranched, laminar glands dense in 2 rows beneath and scattered
above. Inflorescence(\ )3-l 5(-32)-flowered, without accessory flow-
ers, often with l-3(-7)-flowered dichasia from up to 6 nodes below,
and sometimes with 1-2 pairs of flowering branches, the whole
narrowly to broadly cylindric or very rarely obpyramidal; pedicels
0.5-1 mm long; bracts foliar. Flowers 10-18 mm in diam.; buds
ovoid, acutely acuminate. Sepals 5, (3.5-)4-6.5(-7) x 0.4-0.8 mm
in diam.; unequal to subequal, linear-subulate, acute, with margin
revolute, 1 -veined, midrib unbranched. Petals 5, pure yellow, spread-
ing, (5-)6-10 x 3-6 mm, c. 1.5 x sepals, obovate to elliptic-
oblanceolate, with apiculus lateral, acute. Stamens c. 50-80(-1 15),
longest 4.5-6.5(-7) mm,c. 0.65-0.8 x petals. Ovary(2)3(4)-merous,
3^4.5 x 0.5-1.2 mm, very narrowly pyramidal-ovoid (almost cylin-
dric), acute, placentation parietal; styles 3, 2-3.5 mm long, c. 0.7 x
ovary, not separating in fruit. Capsule (4.5-)5-7 x (1.4-)2-3 mm,
very narrowly conic to cylindric. Seeds reddish brown, c. 0.5 mm
long, scarcely carinate; testa finely reticulate. 2n = 1 8 (n = 9, Adams
in Robson & Adams, 1 968, for H. exile).

In moist habitats (open stream banks, pond margins and ditches),
also (in Cuba) in dry meadows and on white sand; lowland (U.S.A.)
to 2000 m (Cuba).

South-eastern U.S.A., mainly in Georgia and Florida but with
outlying stations in Alabama, South Carolina and North Carolina;
also in Cuba (including Isla de Pinos) and Belize.

When considered over its whole range, H. nitidum can be divided
into three subspecies: a) Cuba and Belize (subsp. cubense), giving
rise to 1 1 .H. limosum in Cuba; b) mainland U.S.A. (subsp. nitidum);
and c) western Cuba and north-western Florida (subsp. exile), giving
rise to 12. H. brachyphyllum. The whole complex seems to have
been derived from a narrow-leaved form of 2. H. prolificum (e.g.
some Arkansas populations), to which subsp. cubense approaches
most closely, and to form a sister-group of the H. fasciculatum
complex (Spp. 12-14) (see Fig. 1, p. 77).

9a Hypericum nitidum subsp. cubense (Turcz.) N. Robson in

Bull. nat. Hist. Mus. Lond. (Bot.) 23: 67 (1993).
Fig. 16A.

H. cubense Turcz. in Bull. Soc. Nat. Moscou 31: 384 (1858). Type:

STUDIES IN HYPERICUM

107

Fig. 16 A. H. nitidum subsp. cubense: (a) habit; (b) leaf, lower surface and T.S.; (c) sepal; (d) petal; (e) ovary; (0 capsule. B. H. fasciculatum: (g) leaf,
lower surface and T.S.; (h) fruit. C. H. lissophloeus: (i) habit. D. H. brachyphyllum: (j) leaf; (k) stem node and leaf bases; (1) capsule. E. H. tenuifolium:
(m) stem node and leaf bases (a, i x !/2; b-m x 4, except b\ g' x 8). A. Whitefoord 1934. B. Judd 2649. C. Godfrey 73983. D. Godfrey 57862. E. Herb.
Biltmore 2563a.

108

N.K.B. ROBSON

Cuba, Oriente Prov., St.-Jago, in cacumine Sierra Moestii, 1500
m, Linden 1692 (KW-holotype; BM!, G!, K!, P!, W!-isotypes).

H. aspalathoides sensu Jennings in Ann. Carneg. Mus. 11: 189
(1919).

H.fasciculatum sensu Alain in Leon & Alain, Fl. Cuba 3: 3 17 (1953)
pro parte, quoad syn. H. cubense et prov. cit. Oriente et Isla de
Pinos; Standley & Williams in Fieldiana (Bot.) 7: 49 (1961);
Lippold in Wiss. Z. Friedr.-Schiller Univ. Jena (Math.-Nat. R.) 19:
380, f. 3 (1970) pro parte, quoad syn. H. cubense et prov. cit.
Oriente et Las Villas et Isla de Pinos.

Shrub or small tree to 4.5 m tall. Leaves (6-)10-25 x 0.5-1.6 mm,
coriaceous, apex rounded-apiculate to rounded, margin tightly
inrolled, usually only midrib visible beneath when dry. Inflores-
cence with terminal dichasium c. 7-20-flowered, sometimes with
paired flowers or few-flowered dichasia from up to 2 nodes below.
Sepals obtuse to rounded-apiculate. Petals (6.5-)8-10 x 5-6.5 mm.
Capsule cylindric to rarely ovoid-conic.

Cuba (Oriente, Las Villas, Isla de Pinos), Belize (El Cayo).

CUBA. Oriente: Sierra de Nipe, Rfo Piloto, 20 April 1919 (fl), Ekman
95 10 (F, K, S, US); Sierra Maestra, Rio Alcarraza, July 1946 (fl), Clemente
5069 (GH, US); Moa, between Cerro Miraflores and Moa, c. 30 km E. of
Sagrade Tanamo, 20 July 1951 (fl), Webster 3892 (GH, MICH, US). Las
Villas: Trinidad Mountains, BuenosAires, 22 June 1941, /toward 5201 (BM,
C, F, G, MO, NY, P, S, SING, U, W, WIS). Isla de Pinos: vicinity of Los
Indies, 13 February 1916 (fl), N. & E. Britton & Wilson 14181 (F, NY, US);
near San Pedro, 8 February 1956 (fl), Morton 10040 (BM, US).

BELIZE. Cayo: Mountain Pine Ridge, Augustine, 450 m, 1 January
1959 (fl & fr), tfwnr 68 (BM, F, MO, NY*, US); Mountain Pine Ridge, Baldy
Beacon and vicinity, 900-1020 m, 22 March 1987 (fl), Davidse & Brunt
33063 (BM, MO*).

9b. Hypericum nitidum subsp. nitidum
Map 6.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 363, f.
174(1988).

Shrub to 3 m or more tall, bushy, many-branched. Leaves 10-18 x
0.5-1.5 mm, subcoriaceous, apex obtuse to rounded-apiculate, mar-
gin relatively loosely inrolled, part of lower surface besides midrib
usually visible. Inflorescence mostly cylindric, with terminal
dichasium 3-15-flowered and lateral dichasia from 2-6 nodes be-
low. Sepals acute to shortly apiculate. Petals 5-7 (-9) x 3-4(-6) mm.
Capsule cylindric.

Southeastern U.S.A., from northern Florida and adjacent Alabama
to southern N. Carolina.

U.S.A. Alabama: Baldwin Co., fide Adams (1962: 26). Florida: Bay Co.,
just E. of Callaway, 5 November 1959 (fr), Adams 350 (DUKE*, F*, FLAS*,
GA*, K, MO*, NSCC*, NY*, SMU*, US*); Walton Co., Seven Runs Creek
on Fla 81. f. 8 km S. of Redbay, 27 January 1962 (fr), Ward & Myint 2876
(BM). Georgia: Brantley Co., c. 30.5 km E. of Waycross, 1.6 km N. of High
Bluff Church, 29 August 1960 (fr), Kuns 246 (WIS); Echols Co., 104 km W.
ofStatesville, lOJune 1961 (st), Adams 818 (K). North Carolina: Brunswick
Co.,fide Adams (1 962: 26). South Carolina: Bamberg Co., US 301 , 0.5 kmN.
of junction with SC 64, 9 August 1967 (fr), Bozeman, Radford & Radford
1 1394 (BM); Lexington Co., Black Creek, 9.6 km W. of Pelion, 8 August
1939 (fr), Godfrey & Try on 1303 (BM, SING).

9c. Hypericum nitidum subsp. exile (W.P. Adams) N. Robson in

Bull. nat. Hist. Mus. Land. (Bot.) 23: 67 (1993).
Map 6.

H. galioides van cubenseGriseb., Cat. PL Cuba: 39 ( 1 866); Jennings
in Ann. Carneg. Mus. 11: 189 (1917); non H. cubense Turcz.
( 1858). Type: Cuba, Pinar del Rio, 24 July 1860-64, Wright2\26

(GOET-holotype; BM!, GH!, NY!-isotypes).

H. galioides var. axillare sensu Griseb., Cat. PI. Cub.: 39 ( 1 866) pro
parte, quoad spec. cit. Wright 2123.

H. galioides sensu Sauvalle, Fl. cub.: 8 (1868).

H.fasciculatum sensu Alain in Leon &Alain, Fl. Cuba 3: 3 1 7 ( 1 953)
pro parte, excl. typum; Lippold in Wiss. Friedr. Schiller Univ. Jena
(Math.-Nat. R.) 19: 380, f. 3 (1970) pro parte, quoad syn. Griseb.
et spec. cit. ex Prov. Pinar del Rio.

H. exile W.P. Adams in Contr. Gray Herb. Harv. no. 189: 33 (1962),
in J. Elisha Mitchell scient. Soc. 89: 70 (1973); Godfrey &
Woolen, Aquatic & wetland pis s.e. U.S., Dicots: 345 (1981);
Clewell, Guide vase, pis Florida Panhandle: 370 ( 1 985); Godfrey,
Trees, shrubs & woody vines n. Florida, etc.: 362, f. 173 (1988).
Type: U.S.A., Florida, Gulf Co., 4 km E. of Port St. Joe, 20 May
1960 (fl), Adams 456 (GH!-holotype; DUKE*, FLAS*, FSU*,
GA*, K!, MO*, NCSC*, NCU*, NY*, SMU*, US!-isotypes).

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 362, f.
173(1988).

Shrub to c. 1 m tall, slender, little-branched. Leaves 9-26 x 0.5-0.8
mm, chartaceous, apex acute to long-acuminate, margin tightly
inrolled, midrib visible beneath. Inflorescence mostly cylindric,
with terminal dichasium 3-7-flowered and lateral dichasia from 3-
6 nodes below. Sepals acute to long-acuminate. Petals 6-7 x 3-4
mm. Capsule cylindric to narrowly conic.

Sandy soil in open pinewoods.

U.S.A. (NW Florida), Cuba (Pinar del Rio, Isla del Pinos).

U.S.A. Florida: Franklin Co., 16 km W. of Apalachicola, Adams 473
(FSU*); Gulf Co., N. of Apalachicola, 5 May 1930 (fl), Moldenke \ 146 (K);
Liberty Co., Apalachicola National Forest, c. 1 .6 km E. of Sumatra, 26 April
1975 (fl), Godfrey 74289 (BM, FSU*).

CUBA. Pinar del Rio: Rio Guao, 26-27 February 1 9 1 1 (fl & fr), N. & E.
Britton & Cowell 9618 (F, GH, K, MO, NY); Herredura, 13 April 1920 (fl),
Ekman 10793 (G, K, S);Vinales, April 1930(fl), Leorc 14329 (GH, US). Isla
de Pinos: fide Jennings (1917) under H. galioides var. cubense.

Although subsp. exile is more variable in Cuba than in Florida, the
variation in the two areas overlaps. There can be therefore little
doubt that the two populations belong to the same taxon.

1 0. Hypericum limosum Griseb., Cat. PI. Cub. : 39 ( 1 866); Lippold
in Wiss. Z. Friedr.-Schiller Univ. Jena (Math.-Nat. R.) 19: 381
(1970) pro parte, excl. syn. H. brachyphyllum pro parte. Type:
Cuba, Pinar del Rio, in terra spongiosa ad Lagunas, pr.[ope] S.
Julian, 1861-1864(11), Wright 2 125 (GOET-holotype; BM!,G!,
GH!, MO!, NY!, P!, S!, US!-isotypes).

Map 6.

Shrublet (or annual herbl) 0. 1 5-0.6 m, erect, branching above base,
with branches slender, lower erect, upper divergent. Stems reddish?,

STUDIES IN HYPER1CUM

109

4-lined and ancipitous when young, soon 2-lined, eventually terete;
cortex exfoliating in strips; bark smooth, thin. Leaves sessile, 3-6 x
0.5-1.1 mm, with those in axillary clusters usually as long, linear-
subulate, with margin revolute, sometimes completely concealing
all but the midrib beneath, the non-midrib area rugose-papillose,
chartaceous, deciduous at basal articulation, apex acute to rounded-
subapiculate, base parallel or narrowly cuneate; midrib unbranched;
laminar glands dense, in 2 rows beneath and towards margin above.
Inflorescence 3-c. 15-flowered, (always?) mixed dichasial/pseudo-
dichotomous, without lower axillary dichasia, the whole diffusely
obconic; pedicels to 0.5 mm long; bracts foliar. Flowers c. 10 mm in
diam.; buds broadly ellipsoid, subacute. Sepals 5, 2.5-3.5 x 0.7-0.8
mm, subequal, linear-oblong to oblanceolate, apiculate-obtuse, with
margin revolute, 1 -veined, midrib unbranched. Petals 5, pure? yel-
low, ascending to spreading, 6-8 x 3.5-5 mm, c. 2.4 x sepals,
curved-obovate, with apiculus lateral, minute or absent. Stamens c.
30, longest 3.5-4 mm, c. 0.5-0.6 x petals. Ovary 3-merous, 2-2.5 x
1 mm, very narrowly pyramidal-ovoid, acute, placentation parietal;
styles 3, 1 .2-1 .5 mm long, c. 0.6 x ovary, separating in fruit. Capsule
4-5 x 2-2.5 mm, cylindric. Seeds not seen.

Margins of lakes, marshes and streams on poor sandy soils; lowland.

Cuba (extreme western Pinar del Rio).

CUBA. Pinar del Rio: Between Guane and Remates, Laguna de Cabo,
near sea level, 23 December 1937 (fl), Killip 3233 (US); Laguna Jovero to
Laguna Herradura, 12 December 191 1 (fl & fr), Shafer 10928 (F, MO, NY,
US); RioGuao, 26-27 February 191 1 (f\),N. & E. Britton & C0W//9618 (U,
US).

H. limosum is, both morphologically and geographically, clearly
derived from the Cuban population of H. nitidum subsp. cubense,
differing from it essentially in habit and by the constantly smaller
leaves, flowers and fruit and mixed inflorescence. H. brachyphyllum
appears to be a parallel derivation from H. nitidum subsp. exile, and
so it is not surprising that it was confused with//, limosum by Adams
(1962: 30) and Lippold (1970).

1 1 . Hypericum brachyphyllum (Spach) Steud., Nomencl. 2nd ed.
1: 787 (1840); W.P. Adams in Contr. Gray Herb. Harv. no. 189:
27 (1962), in J. Elisha Mitchell scient. Soc. 87: 70 (1973); R.
Long & Lakela, Fl. Trap. Florida: 608 (1971); R.C. Clark in
Ann. Mo. hot. Gdn 58: 209 (1971); Godfrey & Woolen, Aquatic
& wetland pis S.W., Dicots: 343 ( 198 1 ); Clewell, Guide vase, pis
Florida Panhandle: 370 ( 1 985); Godfrey, Trees, shrubs & woody
vines n. Florida, etc.: 357, f. 170 (1988). Type as for Myriandra
brachyphyllum Spach.

Fig. 16D, Map 7.

H. aspalathoides sensu Elliott, Sketch hot. S. Carolina 2: 27 ( 1 821 );
Chapm., Fl. South. U.S. 3rd ed.: 57 (1897).

Myriandra brachyphylla Spach, Hist. nat. veg. Phan. 5: 435 (1836),
in Ann. Sci. nat. (Bot.) II, 5: 365 (1836); K. Koch, Hon. dendrol.:
66 ( 1 853) pro parte, excl. syn.Type: U.S.A., Florida, Apalachicola,
1835 (fr), Drummond s.n. (P!-holotype; GH!, K!, W!-isotypes).

H. fasciculatum var. aspalathoides sensu Chapm., Fl. South. U.S.:
40(1865).

H. fasciculatum sensu Coulter in Bot. Gaz. 11: 85 (1886) pro parte,
quoad syn. Myriandra brachyphylla.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 357, f.
170(1988).

Map 7 Sect. 20: 1 1 H. brachyphyllum • specimens, O records
(incomplete, as Adams (1959) did not differentiate Spp. 9 and 13).

Shrub (0.3-)0.5-1(-1.5) m tall, usually 1-stemmed, erect, forming
rounded bush, with branches erect Stems reddish to brownish,
sometimes 4-lined when very young (especially just below node),
very soon 2-lined and ancipitous, not becoming terete; cortex exfo-
liating in strips or plates; bark smooth, thin. Leaves sessile, 6-1 1 (-12)
x 0.5-0.7 mm, with those in axillary clusters usually half as long to
as long, linear, plano-convex, glossy green above, with margin
revolute but not concealing all surface beneath, the non-rib area
almost smooth, chartaceous, deciduous at basal articulation (midrib
base not prominent, cf. 9. H. tenuifolium), apex rounded-apiculate,
base parallel; midrib unbranched; laminar glands few, dense, in 2
rows beneath. Inflorescence 3-c. 15-flowered, dichasial or occa-
sionally mixed dichasial/pseudo-dichotomous, without accessory
flowers, with 3-5-flowered dichasia or flowering branches from up
to c. 10 nodes below, the whole ± narrowly cylindric; pedicel absent
or up to c. 1 mm; bracts foliar. Flowers 10-13 mm in diam.; buds
narrowly ovoid-conic. Sepals 5, 2.5-4.5 x 0.5-1 mm, unequal,
linear, acute, 1 -veined. Petals 5, bright yellow, spreading, 5-8 x 2.5-
5 mm, obovate-spathulate, with apiculus lateral, acute. Stamens c.
40-45, longest 4-6 mm,c. 0.75-0.8 x petals. Ovary 3-merous, 2-2.5
x 0.5 mm, very narrowly cylindric, acute, placentation parietal;
styles 3, 3 mm long, 1.2-1.5 x ovary, separating or deciduous in
fruit. Capsule 3.5-5 x 1.5-2 mm, narrowly cylindric to narrowly
ovoid-conic, exceeding sepals, thinly coriaceous. Seeds dull brown,
0.4-0.6 mm long; testa finely reticulate (alveoli circular-hexagonal).
2n = 18 (n = 9) (Adams in Robson & Adams, 1968).

In moist habitats (pine flatwoods, pond margins, borrow pits, swamp
woodland); lowland.

U.S.A. (Georgia and Florida to Louisiana).

110

N.K.B. ROBSON

U.S.A. Alabama: Geneva Co., by county 4, c. 16 km E. of Florala, 2
August 1971 (fl), Krai 43455 (BM); *Covington Co., c. 1 1.8 km S. of Opp,
Skinners 27453 (FSU). Florida: Bay Co., West Bay, 1 2 August 1954 (fl), Ford
4526 (BM). Collier Co., Deep Lake Strand, c. 8 km E. of Miles City, 20
September 1965 (fr). Ward 5233 (BM); Liberty Co., 0.16 km N. of junction
Fla 20 and Fla 276, 14 November 1959 (fr), Adams 372 (K, MO); Wakulla
Co., 1.6 km N. of St. Marks, 17 October 1958 (fr), Godfrey 57862 (BM,
FSU*). Georgia: *Coffee Co., 13 km E. of Douglas, Adams 830 (FSU, GA);
*Early Co., 14.2 km SE of Blakeley, Adams 790 (DUKE, FSU, GA, USF).
Louisiana: New Orleans Par, New Orleans, 1832 (fl), Drummond 51 (BM);
Vernon Par., Anacoco, 60 m, 14 July 1964 (fl), Demaree 50849 (BM).
Mississippi: *Forrest Co., Rte 13, SW of Young, 2 August 1987 (fl). Hill
18458 (MO): Jackson Co., Ocean Springs, 24 August 1951 (fl), Demaree
31293(BM, H).

H. brachyphyllum appears to be a derivative of 1 Oc. H nitidum subsp.
exile. It is very similar to the more reduced form of that subspecies
in western Cuba, but less so to the larger (type) form in Cuba and
north-western Florida. It can be distinguished from subsp. exile by
the bushier habit, usually smaller leaves and smaller flowers, shorter
styles and smaller capsule. It can, on the other hand, be confused
superficially with 9. H. tenuifolium (a reduced 'form' of//, galioides,
not of H. nitidum), but it has a bushy habit, 2-sided shoots, glossy
leaves without prominent base or subapical hydathode, and finely
reticulate seeds, and it grows in wet habitats.

12. Hypericum lissophloeus W.P. Adams in Contr. Gray Herb.
Han', no. 189: 21 (1962), in 7. Elisha Mitchell sclent. Soc. 89: 70
(1973); Ward & Godfrey in D. Ward, Rare & endangered biota
of Florida, 5. Plants: 35 (1980); Godfrey & Woolen, Aquatic &
wetland pis. s.e. U.S. Dicots: 343 (1981); Clewell, Guide vase,
pis Florida Panhandle: 371 (1985); Godfrey, Trees, shrubs &
woody vines n. Florida, etc.: 360, f. 172 (1988). Type: U.S.A,
Florida, Bay Co., shores of Merial Lake, c. 16 km N. of Panama
City, 14 June 1960 (fl & fr), Godfrey & Triplet! 59844 (GH-
holotype; DUKE, F!, FLAS, FSU, GA, IA, ILL, K!, MSC, NY,
SMU, US!-isotypes).

Fig. 16C, Map 3.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 360, f.
172(1988).

Shrub to 4 m tall, erect, slender, sparsely branched, forming dense
clumps sometimes with prop-roots, with branches suberect to as-
cending. Sterns silvery-brown (gun-metal colour), light reddish
brown beneath, 4- lined, ancipitous and glaucous when young, soon
4-angled, eventually terete; cortex exfoliating in large thin curled

plates; bark chestnut-brown, smooth, thin, polished, becoming me-
tallic-silvery. Leaves sessile, (9-) 12- 17 x 0.5-0.75 mm, with those
in axillary clusters almost as long, linear-subulate to acicular,
incurved, with revolute margin not wholly obscuring papillose lower
surface, glaucous when young, subcoriaceous, deciduous at basal
articulation, apex obtuse to rounded, base not or scarcely expanded;
midrib unbranched; laminar glands numerous, especially visible
beneath, marginal glands dense. Inflorescence 1-3-flowered, usu-
ally with paired flowers or triads from up to 9 nodes below, the whole
very narrowly cylindric; pedicels 2-3 mm long; bracts foliar. Flow-
ers c. 20 mm in diam.; buds ovoid, acute. Sepals 5, 7-8 x 0.5-0.75
mm, subequal, linear-subulate, acute, 1 -veined, deciduous. Petals 5,
bright yellow, spreading?, 10-12 x 5-6 mm, obovate-spathulate,
with apiculus lateral, acute, rather short. Stamens 170-221, longest
8-9 mm, 0.75-0.8 x petals. Ovary 3(4)-merous, c. 3.5 x 1.2 mm,
narrowly ellipsoid, placentation parietal; styles 3(4), c. 5 mm long,
c. 1.4 x ovary, separating in fruit. Capsule 6-7 x 2.5-3.5 mm,
narrowly ovoid to ellipsoid, shorter than sepals, thinly coriaceous.
Seeds tan to dark brown, 1-1.6 mm long, shallowly carinate; testa
very coarsely reticulate-sulcate. 2n = 18, n = 9 (Lewis, Stripling &
Ross, 1962).

In sandy soil on shores of sinkhole ponds and lakes, often in water to
1.5 m deep; lowland.

U.S.A. (NW Florida, Bay and Washington Counties only).

U.S.A. Florida: Bay Co., Lake Merial, E. of Fla 77, 24 km N. of Panama
City, 23 August 1966 (fr), Ward 5958 (BM, FSU*); Washington Co., Long
Pond, 7 km S. of New Hope, Fla 77, 12 October 1974 (fr), Godfrey 73983
(BM, FLAS*).

As Adams (1962) remarked, H. lissophloeus is distinguished from
the H.fasciculatum complex (Spp. 12-14) by many features, includ-
ing the smooth polished metallic bark (which exfoliates like a
species of Betula), the slender, wand-like, lax or drooping younger
stems, the large seeds with furrowed testa, and the glaucous young
shoots; and it sometimes grows in association with lOb. H. nitidum
subsp. nitidum and 13. H. fasciculatum, remaining distinct from
both. Adams failed to mention the large capsules and wholly 1-3-
flowered lateral inflorescence-branches, which help to remove H.
lissophloeus from both the H. nitidum and the H. fasciculatum
groups, despite the linear-subulate leaves. Indeed, the capsules are
nearer in form to those of H. prolificum than to those of H.
fasciculatum and even more different from the narrow capsules of
the H. nitidum group. It seems most appropriate, therefore, to regard
H. lissophloeus as an early development of the evolutionary line
from H. prolificum to H. fasciculatum, after the departure of the H.
nitidum complex.

13. Hypericum fasciculatum Lam., Encycl. 4: 160 (1797); Pursh,
Fl. Amer. sept. 2: 376 (1814); Elliott, Sketch hot. S. Carolina 2:
28 (1821); Choisy, Prodr. monogr. Hyperic.: 59 (1821), in DC.,
Prodr. 1: 554 ( 1 824); Torrey & Gray,V/. N. Amer. 1: 160 (1838)
pro parte, excl. [var.] p axillare et syn. H. tenuifolium Pursh et//.
coris? Walter et H. michauxii Poiret; Trev., Hyperic. animadv.:
1 5 ( 1 86 1 ) pro parte quoad typum; Chapman, Fl. South. U.S. : 40
(1865) excl. var. aspalathoides; Coulter in Bot. Gaz. 11: 85
( 1 886) pro parte excl. syn. H. nitidum Lam. et Myriandra spp.,
in A. Gray, Syn. Fl. N. Amer. 1: 286 ( 1 897); R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 2nd ed. 21: 1 80 ( 1 925) pro parte, excl.
syn. H. nitidum Lam.; Small, Man. s.e.fl.: 872 (1933); Lott in/.
Arnold Arbor. 19: 150 (1938) pro parte, quoad typum et syn.
Myriandra brathydis Spach; R. A. Vines, Trees, shrubs & woody
vines of S.W.: 757 (1960); W.P. Adams in Contr. Gray Herb.
Harv. no. 1 89: 24 ( 1 962), in/ Elisha Mitchell sclent. Soc. 89: 70

STUDIES IN HYPERICUM

111

Map 8 Sect. 20: 13. H. fasciculatum • specimens, O records
(incomplete, as Adams (1959) did not differentiate Spp. 9 and 13).

(1973); Radford, Ahles & Bell, Man. vase. fl. Carolina*: 712
(1968); Long & Lakela, Fl. trop. Florida: 607 (1971); R.C.
Clark in Ann. Mo. hot. Gdn 58: 209 ( 1 97 1 ); Godfrey & Wooten,
Aquatic & wetland pis s.e. U.S. Dicots.: 245 (1981); Clewell,
Guide vase, pis Florida Panhandle: 37 1 (1985); Godfrey, Trees,
shrubs & woody vines n. Florida, etc.: 364, f. 175 (1988); N.
Robson in Cullen et al., Eur. Gdn Fl. 4: 69 (1995). Type: U.S.A.
S. Carolina, 'Carolina' (fl), Fraser s.n. (P-LA-holotype, BM!-
microfiche, GH-photograph; BM!, G-DC!, K!-isotypes). The
Lamarck specimen is labelled merely 'Carolina', whereas the
BM specimen has 'Carolina australis' and the Geneva and Kew
specimens 'Carolina med.' ; but they may all be duplicates of one
Fraser collection.
Fig. 16B, Map 8.

H. aspalathoides Willd., Sp. pi. 3: 1451 (1802); Pursh, Fl. Amer.

sept 2: 376 (1814), nom. illegit. (Art. 63). Type as for H.

fasciculatum Lam.
IH.fulgidum Raf., Fl. ludov.: 88 (1817). Type: U.S.A., Louisiana?,

Robin (?). Adams (1962) suggested that this epithet might apply

to H. fasciculatum, despite the absence of that species today from

Louisiana. I have not seen the type, but the description could also

apply to either of the small ('2ft') linear-leaved members of sect.

Myriandra that do occur in that state, viz. H. galioides and H.

brachyphyllum.
Myriandra brathydis Spach, Hist. nat. veg., Phan. 5: 436 (1836),

nom. illegit. (Art. 63). Type as for Hypericum fasciculatum Lam.
Hype ricum fasciculatum var. aspalathoides (Willd.) Torrey & Gray,

Fl. N. Amer. 1: 672 ( 1 840) pro parte, quoad basionym; S. Watson,

Bibliogr. index N. Amer. hot. 1: 126 (1878).
IBrathydiumfuJgidum (Raf.) K. Koch, Hort. dendrol.: 67 (1853).
Hypericum galioides var. fasciculatum (Lam.) Svenson in Rhodora

42: 12, t. 587 f. 1 (1940).

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 364, f.

175(1988).

Shrub to 1 .5(-3) m tall, erect, much branched above but not tree-like,
with branches erect or narrowly ascending. Stems orange-brown, 6-
lined and strongly ancipitous when young, soon narrowly 2-winged,
eventually terete; cortex exfoliating in thin papery sheets or plates

exposing red bark beneath; bark either corky or spongy with incon-
spicuous laticifers, thick. Leaves dark green (cf. H. chapmanii),
sessile, (8-)10-17(-20) x 0.7-1 mm, with those in axils as long,
linear-subulate, sometimes slightly broadened distally, with margin
revolute, overarching all but the ± raised midrib area beneath and
forming 2 longitudinal grooves lined with papillae, chartaceous to
subcoriaceous, deciduous at basal articulation, apex rounded-
apiculate to acute, base parallel; midrib unbranched; laminar glands
dense, in 2 often uneven rows beneath and scattered above. Inflores-
cence (3-)7-32-flowered, without accessory flowers, sometimes
with single flowers or 3-5-flowered dichasia from up to 3 nodes
below, the main inflorescence sometimes reverting to vegetative
growth before producing more flowers, the whole rounded-pyrami-
dal to corymbiform; pedicels absent or almost so; bracts foliar.
Flowers 13-16 mm in diam.; buds ovoid, acute. Sepals 5, (3)4.5-8(-
10) x 0.5 mm, unequal, linear-subulate, acute, with margin revolute,
1 -veined, midrib unbranched, eventually deciduous. Petals 5, bright
yellow, 6-9 x 4-5 mm, c. 1.2 x sepals, obovate-spathulate, with
apiculus lateral, acute. Stamens c. 70-100, longest 5-6.5 mm, 0.7-
0.85 x petals. Ovary 3-merous, 2.5-3 x 1-1.5 mm, very narrowly
pyramidal-ovoid, acute, placentation parietal; styles 3, 2.5-3 mm
long, equalling ovary, separating in fruit. Capsule 5.5 x 2.5-3 mm,
ovoid-conic to ovoid-ellipsoid, 3-sulcate. Seeds dull brown, c. 0.4
mm long, ecarinate; testa finely foveolate-reticulate. 2n = 1 8 (n = 9,
Adams in Robson & Adams, 1968).

Margins of cypress ponds and lakes, marshes and ditches; lowland.

U.S.A. (southeastern North Carolina to southern Mississippi, in-
cluding all Florida).

U.S.A. Alabama: Baldwin Co., just N. of junction 1-10 by Ala 59, 9.6
km S. of Stapleton, Hwy 21, 8 June 1971 (fl), Krai 43080 (BM). Florida:
Brevard Co., Okeechobee region, 3 August 1903 (fl & fr), Fredholm5930(K,
MO); Dade Co., Humbugus Prairie, 8 July 1915 (fl & fr), Small. Mosier &
Small 6878 (K, MO); Duval Co.?, near Jacksonville, May 18— (fl), Curtiss
258 (BM, K, FR); Liberty Co., 5.6 km N. of Bristol, 14 November 1959 (fr),
AdamsXA (K, MO); Lake Co., Kirkland Lake, 20 March 1967 (fl), Harriman
1051 (H); Martin Co., N. of St. Lucie R., E. side of US 1, 5 May 1962 (fl &
fr), Lakela 25048 (BM); Orange Co., 6.4 km S. of Oakland, 8 August 1958
(fr), Godfrey 57329 (BM, FSU*); Palm Beach Co., 24 km W. of West Palm
Beach, 18 June 1937 (fl & fr), Brooks & Murray 5 (BM). Georgia: Colquitt
Co., 9.6 km SE of Moultrie, 8 June 1 958 (fl), A dams 1 3 (K, MO); Sumter Co.,
15 July 1901 (fl),//arper!076(BM,MO).*EcholsCo.,c. 1 3 km W. of Fargo,
Adams 820 (FSU, GA, GH). Mississippi: southern Mississippi, fide Adams
(1962: 24). N. Carolina: no precise locality, n.d. (fr), Nuttall s.n. (BM). S.
Carolina: *Lexington Co., 9.6 km S. of Columbia, 7 August 1939 (fr),
Godfrey & Tryon 1201 (MO). Texas: ?

112

N.K.B. ROBSON

The epithetfasciculatum refers to the crowded leaves of the axillary
shoots, which in this species are usually as long or almost as long as
the subtending ones, in contrast to those of species of the//, nitidum
group, where they are usually shorter than the subtending leaves. In
addition, H. fasciculatum differs from H. nitidum in its thicker,
sometimes spongy bark; its leaves with the lower surface raised,
forming two lateral grooves; its inflorescence, which is more
corymbiform than cylindric; and its ovoid-ellipsoid rather than
cylindric capsules. Both species occur (in the United States) from
southern N. Carolina to southern Alabama, but the distribution of//.
fasciculatum extends throughout peninsular Florida and into Missis-
sippi, whereas H. nitidum subsp. nitidum does not occur south of the
Florida panhandle or west of Alabama.

14. Hypericum chapmanii W.P. Adams in Contr. Gray Herb. Harv.
no. 189: 22 (1962); Godfrey & Wooten, Aquatic & wetland pis
s.e. U.S. Dicots: 346 (1981); Clewell, Guide vase, pis Florida
Panhandle: 37 1 ( 1 985); Godfrey, Trees, shrubs & woody vines n.
Florida, etc.: 366, f. 176 (1988). Type as for H. arborescens
Chapm. non Vahl.

Map 3.

H. arborescens Chapm., Fl. South. U.S. 2nd ed., suppl. 2: 680
(1892), non Vahl (1791). Type: Adams (1962) failed to locate a
Chapman specimen of this species dating from 1 892 or earlier,
and I have not seen one either. Herb. Biltmore no. 5735a (see
below) serves as a representative specimen until a type is discov-
ered.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 366, f.
176(1988).

Shrub, usually one-stemmed and tree-like, up to 4 m tall, erect, with
branches narrowly ascending. Stems orange-brown, 4-lined and
ancipitous when young, soon 4-angled, eventually terete; cortex
exfoliating in thin papery sheets or plates, exposing reddish brown to
cinnamon bark beneath; bark spongy, thick, appearing fluted when
torn apart owing to the presence of large vertically aligned laticifers,
furrowing and disintegrating to expose stringy covering of the
laticifers. Leaves light green (cf. H. fasciculatum), sessile, (8-)l 1-
16(-25) x 5-7 mm, with those in axils as long, linear-subulate, with
margin revolute, overarching all but the raised midrib area beneath
and forming 2 longitudinal grooves lined with papillae, chartaceous,
deciduous at basal articulation, apex narrowly acute, base parallel or
slightly expanded; midrib unbranched; laminar glands ± dense, in 2
regular rows beneath and scattered above. Inflorescence 1-3-flow-
ered, without accessory flowers, often with 1-3 flowers in axil of
leaves at 1-2 nodes below, the whole then shortly cylindric; pedicels

absent; bracts foliar. Flowers 12-15 mm indiam.; buds ovoid, acute.
Sepals 5, 5-7 x 0.5 mm, unequal, linear-subulate, acute, with margin
revolute, 1 -veined, midrib unbranched. Petals 5, bright? yellow, 7-
9 x 3^4.5 mm, c. 1 .3-1 .4 x sepals, oblong-spathulate, with apiculus
lateral, acute. Stamens c. 75, longest 5-5.5 mm (or longer?), c. 0.7 x
petals. Ovary 3-merous, c. 3 x 1 mm, very narrowly pyramidal-
ovoid, acute, placentation parietal; styles 3, 2.5-4 mm long, c. 0.7-1
x ovary, slightly separating at apex in fruit. Capsule c. 6 x 2.4 mm,
narrowly pyramidal-ovoid, 3-sulcate. Seeds dull brown, 0.6-0.8 mm
long, ecarinate?; testa finely foveolate-reticulate.

Flatwoods, depressions, margins of cypress ponds, and borrow pits;
lowland.

U.S.A. (NW Florida).

U.S.A. Florida: *Bay Co., NEofYicksburgh,Adams51 3 (FSU); Franklin
Co.?, Apalachicola, near the coast, July-September 1893 (fl), Chapman in
Herb Biltmore 5735a (A*, BM, GH, NY*); Gulf Co., 4 km SE of Port St Joe,
1 8 June 1 958, Adams 45 (K); *Santa Rosa Co., Tyson 485 (FLAS); *Liberty
Co., c. 5 km SW of Kern, c. 8 km SW of Wilma, Apalachicola Nat. For., 9-
12 m, 13 September 1989 (fr), Orzell & Bridges 12061 (MO).

H. chapmanii is apparently a local derivative of H. fasciculatum,
differing from that species in its taller single-stemmed habit, thicker
stems (100-150 mm as opposed to up to 50 mm) that are less
markedly ancipitous when young, spongy bark with large laticifers
that give a striated or fluted aspect and darken with age, lighter green
leaves that are markedly ascending, and a fewer-flowered inflores-
cence. It would be interesting to know how these differences arose
and are apparently maintained between two species with similar
habitats and with the distributional area of one completely within
that of the other.

Subsect. 2. Pseudobrathydium R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 3(6): 214 (1893) sub sect.
Brathydium; W.P. Adams in Contr. Gray Herb. Harv. no.
189: 33 (1962) emend. Type: H. buckleyi M.A. Curtis
(holotype).

Dwarf wiry shrubs with leaves not articulated at base, persistent or
deciduous above base; inflorescence-branching dichasial or absent,
acropetal; sepals 5, subequal, persistent; stamens c. 100, persistent;
petals 5; styles and placentae 3, placentation incompletely axile.
Species no. 15.

15. Hypericum buckleyi M.A. Curtis in Amer. J. Sci.44: 80(1843)
['buckleii']; Chapm., Fl. South. U.S.: 39 (1865); S. Watson,
Bibliogr. index N. Amer. hot.: 125 (1878); Coulter in Bot. Gaz.
11: 83 (1886), inA. Gray, Syn.fl. N.Amer. 1:285 (1897); Sargent
in Gdn Forest 4: 581, f. 10 (1891); C.K. Schneider, ///. Handb.
Laubholzk. 2: 33, f. 222a (1912); R. Keller in Engl. & Prantl,
Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Small, Man. s.e.fl:
873 (1933): Svenson in Rhodora 42: 19 (1940); Rehd., Man.
cult, trees 2nd ed.: 641 ( 1940); W.P. Adams in Contr. Gray Herb.
Harv. no. 189: 37 (1962), in 7. Elisha Mitchell scient. Soc. 89: 70
(1973); Radford, Ahles & Bell, Man. vase, pis Carolinas: 713
(1968); Bean, Trees & shrubs hardy in Br. Isles 8th ed. 2: 414
(1973); Godfrey & Wooten, Aquatic & wetland pis s.e. U.S.
Dicots.: 348 (1981); Barker & Cheek in Kew Mag. 11: 65-69, t.
245 ( 1 994); Wilbur inCastanea 60: 1 66-1 67 ( 1 995) ['buckleu];
N. Robson in Cullen et al., Eur. Gdn Fl. 4: 69 (1995) [sphalm.
'buckleiyi'].Type: U. S. A. , N. Carolina, Macon Co., 'inmontibus
Carolinae et Georgiae, Pickens Nose', June [1842] (fl), Buckley
s.n. (GH-lectotype, P. Adams, 1962; BM!, NY-syntypes).

Map 5.

STUDIES IN HYPERICUM

Icones: Sargent in Gdn Forest 4: 581, f. 10 (1891); Justice & Bell,
Wild/Is N. Carolina: 1 18 (1968); Barker & Cheek in Kew Mag. 11:
1.245(1994).

Shrub 0.05-0.45 m tall, decumbent, spreading, with branches erect
to ascending from decumbent rooting base, forming low compact
mats. Stems reddish, 4-lined and ancipitous; cortex exfoliating in
strips; bark reddish brown, smooth, thin. Leaves sessile or with
pseudopetiole to 1 mm long; lamina 4-25 x 2-12 mm, oblong or
elliptic to oblanceolate or obovate, plane, paler beneath, chartaceous,
persistent or breaking off at top of pseudopetiole, apex rounded,
base cuneate; venation obscure: (2)3(4) pairs of main laterals with-
out visible subsidiaries or tertiaries, only midrib prominent; laminar
glands dense. Inflorescence l(3-5)-flowered, terminal; pedicels
1.5-3 mm long; bracts foliar. Flowers 20-25 mm in diam.; buds
broadly ovoid. Sepals 5, 4-5 x 2.5-3 mm (to 6 x 3.5 mm in fruit), not
imbricate, subequal, persistent, broadly elliptic to elliptic-spathulate
or obovate, obtuse, plane, basal veins 3, not or obscurely branched.
Petals 5, golden yellow, becoming reflexed, 6-10.5 x 3-5 mm, 1 .5-
2 x sepals, oblanceolate, with apiculus lateral, obtuse. Stamens c.
100, longest 6-9 mm, c. 0.8 x petals. Ovary 3-merous, 3—4 x 2-3
mm, narrowly ovoid, acute, placentation incompletely axile; styles
3, 2.5^ mm long, 0.85-1 x ovary, separating in fruit. Capsule 8-12
x c. 5 mm, narrowly ovoid to ovoid-cylindric, acute, rounded-
trigonous, exceeding sepals, thinly coriaceous. Seeds 1.5-2 mm
long, dark brown, narrowly to broadly carinate; testa finely
scalariform.

Seepage areas, moist crevices and sometimes ditches and road
embankments; 900-1560 m.

Eastern U.S.A. (southern Appalachian Mts - N. Carolina and Geor-
gia - and in adjacent S. Carolina).

U.S.A. N. Carolina: Haywood Co., Mt Pisgah, 14 August 1972 (fr),
Herb. Biltmore 1319b (BM, G, H, JE, K, MO); Macon Co., Highlands, Mt
Satulah, 25 June 1960 (fl), Adams 528 (K). S. Carolina: Greenville Co.,
French Broad R. near Caesar's Head, n.d. (fr), Gray s.n. (K). Georgia: ? Co.
summit of Thomas Bald, 1560 m, August 1893 (fr), Small s.n. (G, H, MO).

Although//, buckleyi differs markedly in habit and leaf-shape from all
other 5-petalled species in sect. Myriandra, the large ovoid capsules
and relatively broad leaves indicate that its nearest relative is H.
prolificum, of which some montane specimens from N. Carolina (e.g.
Leonard, Radford & Moore 1 805 (BM)) resemble it most closely.

The original spelling, buckleii, was correct until recently accord-
ing to the International Code of Botanical Nomenclature (Art. 73.1),
as Curtis's latinization of Buckley was clearly intentional. There was
therefore no alternative to rejecting the usual spelling, buckleyi, pace
Adams ( 1 962: 37); and I adopted the form buckleii in The New Royal
Horticultural Society Dictionary of Gardening (Robson, 1992),
European Garden Flora (Robson, 1995)5 and various herbarium

5 But my last-minute attempt to correct the spelling resulted in the publication of
another variant: bucklei\i.

113

labels; and Wilbur ( 1 995) also pointed out this necessary change. In
the new (Tokyo) edition of the I.C.B.N. (1994), however, spellings
of epithets based on latinizations of modern names have been
prohibited retrospectively (Art. 60.1 1), so the form buckleyi is now
the obligate one. Barker & Cheek (1994) reached the same conclu-
sion; but their statement that the leaves of H. buckleyi have black
glandular dots is erroneous.

Subsect. 3. Suturosperma R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 3(6): 214 (1893).

Isophyllum Spach, Hist. nat. veg. Phan. 5: 432 (1836), \nAnnls Sci.

nat. (Bot.) II, 5: 367 (1836), non Isophyllum Hoffman (1814).

Type: /. drummondii Spach (= Hypericum microsepalum (Torrey

& Gray) A. Gray ex S. Watson).
Ascyrum b. Isophyllum (Spach) Endl., Gen. pi: 1032 (1840) status

ignot.
Crookea Small, Fl. s.e. U.S.: 786, 1335 (1903). Type: C. microsepala

(Torrey & Gray) Small (= Hypericum microsepalum (Torrey &

Gray) A. Gray ex S. Watson).
Hypericum sect. Isophyllum (Spach) W.P. Adams in Contr. Gray

Herb. Harv. no. 189: 36 (1962), nom. synon.

Shrubs or perennial herbs with leaves not articulated at base,
persistent or deciduous above base; inflorescence-branching
dichasial, mainly acropetal; petals 5 (4-3); stamens 30-95, decidu-
ous (Spp. 1 6, 1 7) or persistent; styles and placentae 3(4), placentation
incompletely axile (Spp. 16, 17) or ± parietal. Species 16-22.

16. Hypericum apocynifolium Small in Bull. Torrey hot. Club 25:
616 (1898), Fl. s.e. U.S.: 788 (1903), Man. s.e.fl.: 871 (1933);
Svenson \nRhodora 42: 15 (1940) pro parte, excl. spec. Harper.;
W.P. Adams in Contr. Gray Herb. Harv. no. 189: 37 (1962);
Correll & Johnson, Man. vase, pis Texas: 1 064 ( 1 970); Godfrey,
Trees, shrubs & woody vines n. Florida, etc.: 371, f. 179e-g
(1988). Type: U.S.A., Arkansas, Miller Co., Texarkana, August
1897 (fl & fr), Heller s.n. (NY-lectotype, Svenson 1940); with-
out precise locality, Leavenworth s.n. (NY-syntype). Small refers
to Texarkana specimens without mentioning Heller, which is
probably why Svenson merely 'suspects' it to be the type. Both
refer to the Leavenworth specimens from Arkansas (H.
nudiflorum var. (3 sensu Torrey & Gray) as additional material.
Svenson (1940) may thus be said to have lectotypified //.
apocynifolium Small; but, in any case, Adams (1962) definitely
states that the Heller specimen is the lectotype.

Map 9.

H. nudiflorum [var.] (3 sensu Torrey & Gray, Fl. N. Amer. 1: 162
(1838).

Map 9 Sect. 20: 16. H. apocynifolium • specimens, D records; 17. H.
nudiflorum • specimens. O records.

114

N.K.B. ROBSON

H. nudiflorum sensu R. A. Vines, Trees, shrubs & woody vines ofS. W. :
754 (1960); W.P. Adams in J. Elisha Mitchell sclent. Soc. 89: 70
(1973); Clewell, Guide vase, pis Florida Panhandle: 370 (1985)
pro parte omnes quoad syn. H. apocynifolium; et auct. plur.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 371 , f.
179e-g(1988).

Shrub 0.4-0.7 m tall (or taller?), erect, with branches ascending.
Stems orange-brown, narrowly 4-winged at first, eventual 1 y 2-lined,
cortex exfoliating in strips, exposing red bark; bark turning brown.
Leaves bright green, subsessile or shortly broadly petiolate (to 1
mm), 20-40 x 12-20 mm, oblong to elliptic-oblong, with margin
plane or rarely recurved, paler or ± glaucous beneath, ± thinly
chartaceous, eventually deciduous at lamina base, apex rounded to
retuse, base ± broadly cuneate; venation: c. 6 pairs main laterals
without intermediates, tertiary reticulation obscure; usually only
midrib prominent; laminar glands small, dense. Inflorescence (1)3-
5(-8)-flowered, terminal, without accessory flowers; pedicels 3-10
mm long; bracts 1 .5-3 mm long, triangular-subulate. Flowers c. 1 5
mm in diam.; buds globose. Sepals 5, 3-5 x 1.5-2.3 m, unequal,
tardily deciduous, spathulate to elliptic or ovate, rounded to acute,
margin plane, basal veins (1)3, unbranched. Petals 5, coppery
yellow, 8-10 x 4-6 mm, c. 2 x sepals, oblong, with apiculus lateral,
acute, very short. Stamens c. 60-80, longest 4-6 mm, 0.65-0.75 x
petals, deciduous. Ovary 3-merous, c. 3 x 1.5 mm, ellipsoid, suba-
cute, placentation incompletely axile; styles 3, 1-1.5 mm long,
c. 0.5 x ovary. Capsule 6-15 x 4.5-7(-8) mm, cylindric-conic,
acute, longer than sepals, thickly coriaceous. Seeds dull brown to
blackish, 1 .8-2 mm long, with low ridge; testa finely scalariform-
reticulate.

Stream banks and moist woods; coastal plain and lower Mississippi
valley.

U.S.A. (scattered localities in south-eastern Oklahoma, southern
Arkansas, western Texas and western Louisiana; also (?) in extreme
south-western Georgia and Florida).

U.S.A. Arkansas: *DrewCo.,Monticello,Demaree 16231 (NY); *Miller
Co., 10 June 1898 (fl), Eggert s.n. (MO). Florida: *Gadsden Co., Aspalaga,
Godfrey 53612 (DUKE, FSU, NCSC). Louisiana: Natchitoches Par.,
Natchitoches, 14 June 1915 (e. fr), Palmer%009 (K); De Soto Par., 3.2 kmW.
of Hunter, 10 August 1938 (fr), Correll & Correll 10178 (DUKE*, GH).
Oklahoma: *Choctaw Co., Leavenworth s.n. (NY). Texas: without precise
locality, n.d. (fr), Wright s.n. (K).

According to Adams (1962), H. apocynifolium cannot be distin-
guished vegetatively from 17. H. nudiflorum. The few-flowered
inflorescence, larger flowers with relatively longer sepals, larger,
thicker-walled capsules and seeds with a ridge (not a keel), however,
all seem to be constant distinguishing characters of//, apocynifolium,
and all indicate an evolutionary position between 2. H. prolificum
and 17. H. nudiflorum. I therefore prefer to regard it as a distinct

species, as did Adams at first (although not later - Adams, 1973),
rather than include it in H. nudiflorum. Adams recorded //.
apocynifolium from the Flint River drainage in extreme south-west
Georgia and the Apalachicola River bluffs in Gadsden Co., Florida;
but specimens from those regions that I have seen belong to //.
nudiflorum.

H. apocynifolium is most closely related to a form of//, prolificum
from Tennessee. It and H. nudiflorum are intermediate between that
species and the H. cistifolium group (Spp. 18-21).

17. Hypericum nudiflorum Michx. ex Willd., Sp. pi. 3: 1456
(1802); Michx., Fl. bor.-amer. 2: 78 (1803); Pursh, Fl. Amer.
sept. 2: 375 ( 1 8 1 4); Elliott, Sketch hot. S. Carolina 2:32(1821);
Choisy, Prod, monogr. Hyperic.: 46 (1821), in DC., Prodr. 1:
548 ( 1 824) pro parte excl. syn. Aiton; Hook. & Arn. in Hook. J.
Bot. 1: 199 (1834) incl. var.; Torrey & Gray, Fl. N. Amer. 1: 162
(1838) excl. [var.] p; Darby, Man. Bot. 2: 35 (1841)
['mediflorum']; A. Gray, Man. Bot.: 53 (1848); Chapm., Fl.
South. U.S.: 41 (1865); S. Watson, Bibliogr. index N. Amer. hot.
1: 128 (1878); Coulter in A. Gray, Syn. fl. N. Amer. 1: 287
(1897); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed.
21: 180 (1925); Small, Man. s.e. fl.: 871 (1933); Svenson in
Rhodora 42: 18 (1940); Rehd., Man. cult, trees 2nd ed.: 640
(1940); J.P. Gillespie in Castanea 24: 29 (1958); R.A. Vines,
Trees, shrubs & woody vines ofS. W. : 764 ( 1 960); W.P. Adams in
Contr. Gray Herb. Harv. no. 189: 36(1962), in/ Elisha Mitchell
scient. Soc. 89: 70 (1973) pro parte, excl. syn. H. apocynifolium;
Radford, Ahles & Bell, Man. vase. Fl. Carolinas: 7J3 (1968);
Correll & Johnston, Man. vase, pis Texas: 1064 (1970); R.C.
Clark in Ann. Mo. hot. Gdn 58: 209 (1971); Godfrey & Wooten,
Aquatic & wetland pis s.e. U.S. Dicots: 348 (1981); Clewell,
Guide vase, pis Florida Panhandle: 372 (1985) pro parte, excl.
syn. H. apocynifolium; Godfrey, Trees, shrubs & woody vines n.
Florida, etc.: 371, f. 179 a-d (1988). Type: U.S.A., S. Carolina,
Berkeley Co., 'Goose Creek & Gafnnet?] place', n.d. (fr),
Michaux s.n. (P-holotype, BM! -microfiche; G-photograph).

Fig. 17A, Map 9.

H. ligustrinum Pursh, Fl. Amer. sept. 2: 375 (1814), in synon.
H. nudiflorum [van] (3 ovatum Choisy in DC., Prodr. 1: 548 (1824).

Type: U.S.A., S. Carolina, Car. meridionale, Fraser s.n. (G-DC!).
?//. nudiflorum [var.] y ramosum Choisy in DC., Prodr. 1: 548

(1824). Type not seen; not in Herb. DC.
Myriandra nudiflora (Michx. ex Willd.) Spach, Hist. nat. veg. Phan.

5: 440 (1836), in Ann. Sci. nat. (Bot.) II, 5: 365 (1836).
Brathydium nudiflorum (Michx. ex Willd.) K. Koch, Hort. dendrol.:

67(1853).
Hypericum cistifolium sensu Coulter in Bot. Gaz. 11: 86 (1886) pro

parte excl. typum.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 371, f.
179a-d (1988).

STUDIES IN HYPER/CUM

115

Fig. 17 A. H. nudiflorum: (a) habit; (b) leaf (part); (c) sepal; (d) petal; (e) capsule. B. H. cistifolium: (f) habit; (g) sepal; (h) capsule. C. H. microsepalum:
(i) habit: (j) sepal; (k) capsule (a, f, i x >/2; c x 2; b, d, e, g, h, j, k x 4). A. Herb. Biltmore 1272b. B. Bouffordet al. 23046. C. Chapman s.n.

116

N.K.B. ROBSON

Shrub 0.5-2 m tall, erect, usually loosely branched, with branches
ascending. Stems narrowly 4-winged at first, eventually terete,
subherbaceous above, becoming brown and woody at base; cortex
exfoliating in strips; bark brown. Leaves pale green, subsessile or
shortly and broadly petiolate (to 2 mm long); lamina 30-70 x 7-25
mm, ovate-lanceolate or elliptic to linear-oblong, with margin plane
and very narrowly pellucid, not or rarely slightly glaucous, thinly
chartaceous, deciduous at lamina base, apex obtuse to rounded, base
cuneate to subcordate; venation: c. 6 pairs main laterals and some-
times intermediates, tertiary reticulation obscure; only midrib
prominent; laminar glands dense. Inflorescence 1-c. 45-flowered,
without accessory flowers, sometimes with \-l(-c. 40)-flowered
dichasia from 1-3 nodes below, the whole corymbiform or some-
times rounded-pyramidal; pedicels 1.5-4 mm long; bracts c. 1.5-3
mm long, triangular-subulate. Flowers 15-20 mm in diam.; buds
ellipsoid, rounded. Sepals 5, 2-5 x 1-1 .5 mm, unequal to subequal,
deciduous, oblanceolate-spathulate or oblong-elliptic to narrowly
triangular, obtuse to acute, margins plane, basal veins 3, unbranched.
Petals 5, pale or coppery yellow, 6-8 x 3^ mm, 2.5-3 x sepals,
oblanceolate-oblong to elliptic-oblong, with apiculus lateral, acute,
short. Stamens c. 80, longest 4-5 mm, 0.7-0.8 x petals, deciduous.
Ovary 3(4)-merous, c. 3 x 1 mm, very narrowly ovoid-ellipsoid,
acute, placentation parietal; styles 3(4), c. 3 mm long, about equal-
ling ovary, remaining appressed in fruit. Capsule 3.5-7 x 3-5 mm,
broadly ellipsoid to ovoid-globose, acute. Seeds 1.5-2 mm long,
black, carinate, conspicuously curved (fide Adams, 1959 ); testa ±
scalariform-reticulate. 2n = 18 (n = 9, Hoar & Haertl, 1932).

Stream banks, moist woodland (deciduous and pinelands) and
swamps, on sand; lowland and plateau (to c. 1000 m).

U.S.A. (Virginia to E. Tennessee, south to NW Florida and west to
SE Louisiana and Texas).

U.S.A. Alabama: no precise locality, 1 832 (fl & fr), Drummond s.n. (K);
*Etowah Co., Gadsden, Vasey 385 (F, GH, NY, PH, US). Florida: Wakulla
Co., prope St Marks, June 1843 (fl), Rugel 466 (BM, G, K); Wakulla Co.,
between Bloxham and Sopchoppy, 25 June 1958 (fl), Godfrey 57135 (BM,
FSU*,GH*). Georgia: Clarke Co., SE of Athens, River Bend East road, 28
June 1 979 (f\&h), Jones 23296 (BM,G,GH*); Lee Co., Mill Creek, 13 July
1901 (e. fr), Harper 1073 (BM, MO*). Louisiana: New Orleans Par., New
Orleans, n.d. (fr), 'Hooker' s.n. (probably co\\. Drummond) (K); St. Tammany
Par., Covington, 1 832 (fr), Drummond^ 19 (BM, K). Mississippi: *Stone Co.,
0.4 km S. of McHenry, Diener 378 (MISSA, TULANE). North Carolina:
Burke Co., Table Rock Mtn, 9 August 1909 (fr), Herb. Biltmore 1001 (G);
Chatham Co., 4.8 km S. of Wilsonville, 1 July 1966 (fl), Pence in Radford
44838 (BM, H); Davie Co., near Farmington, 3 July 1 897 (fr), Herb. Biltmore
1272d (JE). South Carolina: Richland Co., Congaree R., 19 June 1855 (fl &
e. fr), Hexamer & Maier s.n. (BM); * Abbeville Co., 8 km SW of Antreville,
Radford 26052 (UNC); see also type. Tennessee: Grundy Co., near Beersheba
Springs, 1 August 1947 (e. fr), Sharp, Shanks & Clebsch 5144 (BM);
*SequatchieCo.,W.ofDunlap,510m, 14 July 1938 (fl & fr),Svenson 9554
(BKL, DUKE, MO, PH, TENN). Virginia: Princess Anne Co., Macon's
Corner, 8 September 1935 (fr), Fernald & Long 4943 (A*, BKL*, GH*, K,
NY*, US*); *Sussex Co., terrace of Nottoway R., c. 5 km NNW of Bethel
Church, 9 September 1946 (fr), Fernald, Long & Clement 15307 (MO).

Correll & Johnston (1970) record H. nudiflorum from eastern Texas
as well as H. apocynifolium; but I have seen no specimens from
there, and Adams (1962) did not include Texas in the distribution of
this species. He likewise failed to record it from Louisiana, although
there are early-nineteenth-century specimens from there, which
suggests that it may now be extinct in that state.

H. nudiflorum is morphologically intermediate between H.
apocynifolium and H. cistifolium. For differentia see these species.

Render (1911) observed pistillody in H. nudiflorum. There were
c. 3-10 sterile structures per flower, situated between pistil and

Map 10 Sect. 20: 18. H. cistifolium • specimens, O records.

stamens and mostly boat-shaped. They differed in size and bore
ovules, rarely except the upper ones. Stamen tissue was often
present, but true anthers were rare.

18. Hypericum cistifolium Lam., Encycl. 4: 158 (1797); Choisy,
Prodr. monogr. Hyperic.: 45 (1821), in DC., Prodr. 1: 547
(1824); Torrey & Gray, Fl. N. Amer. 1: 673 (1840); Chapm., Fl.
South. U. S.: 41 (1865); S. Watson, Bibliogr. index N. Amer. hot.:
125 (1878) pro parte, excl. syn. H. nudiflorum; Coulter in A.
Gray, Syn.fl. N. Amer. 1: 287 (1897); R. Keller in Engl. & Prantl,
Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Svenson in Rhodora
42: 17 (1940); Rehd., Man. cult, trees 2nd ed.: 640 (1940); R.A.
Vines, Trees, shrubs & woody vines of S.W.: 756 (1960); W.P.
Adams in Contr. Gray Herb. Harv. no. 189: 38 (1962), in J.
Elisha Mitchell sclent. Soc. 89: 70 (1973); Radford, Ahles &
Bell, Man. vase. fl. Carolinas: 713 (1968); R.C. Clark in Ann.
Mo. hot. Gdn 58: 209 (1971); R. Long & Lakela, Fl. Trop.
Florida: 607 ( 1 97 1 ); Godfrey & Wooten, Aquatic & wetland pis
s.e. U.S. Dicots.: 348 (1981); Clewell, Guide vase, pis Florida
Panhandle: 371(1 985); Godfrey, Trees, shrubs & woody vines n.
Florida, etc.: 369, f. 178 (1988); N. Robson in Eur. Gdn Fl. 4:
69, ff. 10.6, 10.12 (1995) ['//. cistiflorum']. Type: U.S.A., no
precise locality or collector (P-LA-holotype; GH-photograph).

Fig. 17B, Map 10.

H. rosmarinifolium Lam., Encycl. 4: 159 (1797); Willd., Sp. pi. 3:
1450 (1802); Choisy, Prodr. monogr. Hyperic.: 45 (1821), in
DC., Prodr. 1 : 547 ( 1 824); S. Watson, BM. index N. Amer. hot. I :
456 (1878); Coulter in A. Gray, Syn.fl. N. Amer. 1: 287 (1897)
pro parte, excl. syn. H. sphaerocarpum. Type: U.S.A., Carolina,
Fraser s.n. (P-LA-holotype).

Brathydium hyssopifoliumSpach, Hist. nat. veg. Phan.5: 445 ( 1 836),
in AnnlsSci. nat. (Bot.) II, 5: 365 (1 836); K. Koch, Hort. dendrol.:
67 (1 853), nom. illegit. (Art. 63). Type as for H. cistifolium Lam.

Hypericum opacum Torrey & Gray, Fl. N. Amer. 1: 163 (1838);
Coulter in Bot. Gaz. 11: 87 (1886), in A. Gray, Syn. Fl. N. Amer.
1: 287 (1897); Sargent in Gdn Forest 5: 304 & t. (1892); Small,
Man. s.e.fl.: 871 (1933). Type: U.S.A., Georgia, Mrs Miller s.n.
(GH-syntype?); Georgia, Dr Loomis s.n. (GH-syntype?); Ala-
bama, Dr Gates s.n. (GH-lectotype, selected here).

H. punctulosum Bertol., Misc. Bot. 13: 18, t. 3 f. 2d-e (1853). Type:
U.S.A., Alabama, Dr Gates s.n. (GH-holotype?). Bertolini's col-
lection at BOLO does not contain an appropriate Gates specimen,
judging from the microfiche set.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 369, f.
178(1988).

STUDIES IN HYPERICUM

17

Shruh (or subshrub), c. 0.5-1 .3 m tall, erect, unbranched or with leaf
clusters or short branches from most nodes and sometimes 1-3
ascending branches from lower half of stem. Stems red-brown, 4-
lined, the subfoliar lines more prominent or wing-like, becoming
terete below; cortex exfoliating in strips; bark reddish brown. Leaves
sessile, 15-40 x (2-)4-8(-10) mm, narrowly oblong or narrowly
elliptic-oblong to triangular-lanceolate, with margin recurved,
densely glaucous beneath, thinly coriaceous, deciduous above base,
apex subacute to rounded, base subcordate to cuneate; venation: one
pair of ascending near-basal laterals and c. 4-5 pairs of obscure
laterals, only midrib prominent the rest often invisible externally;
laminar glands small, dense. Inflorescence (7-)15-c. 65-flowered,
of regular dichasia, dense, without accessory flowers, sometimes
with 3-c. 65-flowered dichasia from 1-2 nodes below and short
flowering branches from a further 1 -4 nodes, the whole corymbiform
(often rounded) to cylindric; pedicels 1-3 mm long; bracts 2-10 mm
long, subulate. Flowers 7-12 mm in diam., buds ellipsoid, acute.
Sepals 5, 1-4 x 1-1.8 mm, unequal, persistent, obovate or broadly
elliptic to oblong, rounded, margin plane, basal veins 3(-5?),
unbranched. Petals 5, bright yellow, 5-8 x 2.5-3 mm, 2 x sepals,
oblanceolate, with apiculus lateral, obtuse, short. Stamens c. 30-50,
longest 3.5^.5 mm, 0.6-0.7 x petals, at least some persistent at least
until fruit matures. Ovary 3-merous, 1.5-2.5 x 1-1.2 mm, narrowly
ovoid-conic, obtuse, 3-lobed with valvular depressions, placentation
parietal; styles 3, 1-2 mm, 0.7-0.8 x ovary, remaining appressed in
fruit. Capsule 4-6 x 3-4 mm, ovoid-cylindric to ovoid-subglobose,
obtuse to rounded, 3-lobed by depressions between carpel margins.
Seeds mustard-yellow, 0.5 mm long, ecarinate; testa foveolate-
reticulate to linear-foveolate. 2n = 18 (n = 9, Adams in Robson &
Adams, 1968).

Moist soil in pine flatwoods, bogs, swamp and marsh margins,
ditches and roadside embankments, on sand; lowland.

U.S.A., coastal plain from North Carolina to Louisiana.

U.S.A. Alabama: Baldwin Co.Jufc Clark ( 197 1 ) andAdams ( 1959: 13);
Mobile Co., Mobile, 1 840 (fl), Gray s.n. (K); *Pike Co., Spring Hill, 1 9 1 9 (fl
& fr), Graves s.n. (MO). Florida: Duval Co., near Jacksonville, August (fl &
fr), Curtiss 253 (BM, F*, FR, GH*, K, MO*, PH*, US*); Lake Co., c. 6.4 km
N. of Altoona, 2 August 1960 (fr), Adams 603 (K); Orange Co., near Taft, 1
June 1952 (fl), Schallert4031 (BM, K). Georgia: Brantley Co., c-. 16 km E. of
Waycross, near Highbluff Church, 29 August 1960 (fr), Kuns 261 (WIS);
Pierce Co., Logue & Bozeman 2136 (BM, H). Louisiana: St Tammany Par.,
Covington, 1832 (fl), Drummond 135 (BM, K); *Tangipahoa Par., 3.2 kmW.
of Robert, Correll 10511 (F, GH, NY, PH). Mississippi: *Harrison Co.,
Biloxi, 25 July 1896 (fl & fr), Pollard 1002 (F, GH. MO, NY, US); Jackson
Co., N. of Ocean Springs, 20 July 1952 (fr), Demaree 32427 (BM). North
Carolina: Onslow Co., SE of Dixon, 7.1 km S. of U.S. 17 on N.C. 210, 24
August 1979 (H & e. fr), Boufford 2 1 552 (BM; CM*); Tyrrell Co., near N.C.
94, 0.8 km S. of Kilkenny, 6 August 1959 (fl), Radford 39250 (K). South
Carolina: *Charleston Co., 8 km NW of McClellanville, 4 August 1939 (fl &
fr), Godfrey & Try-on 1114 (DUKE, F, GH, MICH. NY, PH, TENN). Texas:
*Hardin Co., Honey Island, Village Mills. 31.5 m, 25 September 1987 (fr),
Orzell & Bridges 5806 (MO).

Adams (1962), following Torrey & Gray (1838), remarked on the

capsules of H. cistifolium being lobed due to dorsal (not the usual
lateral) compression of the three valves. In this they contrast with the
unlobed globose capsules of H. sphaerocarpum.

Torrey & Gray (1838: 163) originally omitted H. cistifolium,
describing it as a new species, H. opacum Torrey & Gray. Although
they realized their mistake on seeing Lamarck's type of//, cistifolium
and corrected it (Torrey & Gray, 1840: 673), the name H. opacum
remained current in the American literature until Svenson (1940)
explained the situation. As a result of this confusion, Coulter ( 1 886a)
regarded H. cistifolium as a later homonym of//, nudij lorum Michx.
ex Willd.

H. cistifolium is closely related to 18. H. sphaerocarpum, having
a more south-eastern distribution. The species can be distinguished
by leaf size, sepal size and fruit shape.

19. Hypericum microsepalum (Torrey & Gray) A. Gray ex S.
Watson, Bibliogr index N. Amer. hot. 1: 456 (1878); Coulter in
Bot. Gaz. 11: 82 (1886), in A. Gray, Syn. fl. N. Amer. 1: 284
(1897); Adams & Robson in Rhodora 63: 15 (1961); W.P.
Adams in Contr. Gray Herb. Harv. no. 189: 43 (1962), in J.
Elisha Mitchell scient. Soc. 89: 68 (1973); Godfrey & Wooten,
Aquatic & wetland pis s.e. U.S. Dicots: 341 (1981); Clewell,
Guide vase, pis Florida Panhandle: 372 (1985); Godfrey, Trees,
shrubs & woody vines n. Florida, etc.: 352, f. 166 (1988). Type
as for Ascyrum microsepalum Torrey & Gray.

Fig. 17C, Map 11.

Isophyllum drummondii Spach, Hist. nat. veg. Phan. 5: 433 (1836),
in Annls Sci. nat. (Bot.) II, 5: 367 (1836), non Hypericum
drummondii (Grev. & Hook.) Torrey & Gray (1838). Type:
U.S.A., Florida, Franklin Co., prope Apalachicola, 1836 (fl),
Drw/n/no«J8(P-holotype;BM!,GH,K!,W!). Adams (1962: 43)
wrongly stated that Sarothra drummondii Grev. & Hook, was the
basionym of this name. It is in fact the basionym of Hypericum
drummondii (Grev. & Hook.) Torrey & Gray (sect. Brathys).

Ascvrum microsepalum Torrey & Gray, Fl. N. Amer. 1: 157 (1838);
A. Gray, Gen. Amer. bor.\:2\2 (1848); Chapm., Fl. South. U.S.:
39 (1865). Type: U.S.A., Florida, 'Middle Florida', Dr Alexander
(NY-lectotype, selected here); Georgia, without precise locality, ?
(NY-syntype).

Hypericum isophyllum Steud., Nomencl. 2nd ed. 1: 788 (1840),
nom. illegit. (Art. 63).

Brathydium microsepalum (Torrey & Gray) K. Koch, Hon. dendrol:
67(1853).

Map 11 Sect. 20: 19. H. microsepalum • specimens, D records; 20. H.
sphaerocarpum • specimens. O records.

118

N.K.B. ROBSON

Crookea microsepala (Torrey & Gray) Small, Fl. s.e. U.S.: 786,
1335 (1903), Man. s.e.fl.: 868 (1933).

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 352, f.
166(1988).

Shrub 0. 1 5-0.7 m tall (or long), erect or more usually decumbent or
ascending, with branches numerous, straggling or ascending. Stems
reddish brown, 4-lined and ancipitous when young, eventually 2-
lined; cortex exfoliating in strips; bark red-brown. Leaves sessile,
5-10(-15) x 1-3 mm, narrowly oblong or oblanceolate to linear,
with margin recurved to subrevolute, paler beneath, thinly coriaceous,
deciduous above base, apex rounded to obtuse, base rounded to
(usually) cuneate; venation obscure: 1-3 pairs of lateral veins;
laminar glands dense to sparse. Inflorescence 1-3-flowered, without
accessory flowers, sometimes with single flowers or triads or flow-
ering branches from up to 4 nodes below, the whole
rounded-pyramidal; pedicels 5-9 mm long; bracts reduced foliar.
Flowers 15-25 mm in diam., buds ovoid, acute. Sepals (3)4(5), 3-5
x 1-1.4 mm, persistent, subequal or (when 5) equal, oblong or
elliptic to linear, obtuse to acute, margin plane, basal veins 3,
unbranched. Petals (3)4(5), bright yellow, 10-12x6-9 mm, 2-2.5 x
sepals, unequal or (when 5) subequal, when 4 the larger pair obovate
and smaller pair obovate-oblong, with apiculus lateral, obtuse, short.
Stamens 60-70, longest c. 6 mm,c. 0.5 x petals. Ovary 3-merous, 3-
3.5 x 1.2 mm, narrowly ellipsoid, scarcely lobed, placentation
parietal; styles 3.3 mm long, 0.85-1 x ovary, sometimes separating
in fruit. Capsule 6-8 x c. 2.5 mm, cylindric-ellipsoid to cylindric or
narrowly ovoid-conic, acute to obtuse. Seeds dark brown, c. 0.9-1
mm long; testa linear-foveolate. 2n = 18 (Lewis, Stripling & Ross,
1962; n = 9, Adams in Robson & Adams, 1968).

Low pine flatwoods, moist to wet, on sand; lowland.

U.S.A. Northern Florida (Panhandle from Walton to Lafayette coun-
ties) and adjacent Georgia.

U.S. A Florida: Gadsden Co., Lawrence, 15 March 1940, Harper 129
(K); Jefferson Co., N. side of Aucilla R., S. of Lament, 6 April 1960 (fl), Dress
& Hanson in Bailey 2022 (BM, G); Wakulla Co., vicinity of Crawfordsville,
Apalachicola National forest, 27 February 1971 (fl), Godfrey 70178 (H).
Georgia: Atkinson Co.,fideAdams (1962: 43); *Brooks Co., 1 5 April 1 892 (fl
& fr), Lighthipe s.n. (MO); Calhoun Co., fide Adams (1962: 43); *Lowndes
Co., 2.4 km W. of Dasher and c. 1 1 km S. of Valdosta, 20 February 1965 (fl),
Faircloth 1655 (MO).

Although H. microsepalum typically has a tetramerous perianth,
Adams ( 1 962) pointed out that the perianth of some flowers on many
plants may be pentamerous, or a pair of sepals and/or petals may be
partially united (Adams & Robson, 1961). In such flowers the
differences in sepal and petal size are almost absent.//, microsepalum
is thus clearly in a transitional stage between Hypericum proper and
'Ascvrum'; but it does not bear a close resemblance to the other

members of 'Ascyrum'. For example, the pairs of sepals are almost
equal, a feature that induced Spach (1836a, b) to distinguish it as
I sop hy Hum.

Whereas Ascyrum proper (Spp. 25-29) appears to be derived from
1. H. fmndosum, the nearest relative of H. microsepalum would
seem to be 17. H. cistifolium. Indeed, all the characters of H.
microsepalum can be considered as reduced states of those of H.
cistifolium, except the much larger petals. Geographically, the distri-
bution of//, microsepalum falls within that of//, cistifolium; and so
it is apparently a neo-endemic, maintained as a distinct species by
some biological factor, possibly different pollinators attracted by the
different size of flower. Adams's comment that H. microsepalum
flowers earlier than other species (mainly February to late April but
sporadically in May and November) may be relevent in this regard.

20. Hypericum sphaerocarpum Michx., Fl. bor.-amer. 2: 78 ( 1 803);
Pursh, Fl.Amer. sept. 2: 378 (1814); Poir.,Encyd. Suppl.3: 1697
(1814) ['sphaerocarpon']; Choisy, Prodr. Monogr. Hyperic.: 46
( 1 82 1 ), in DC., Prodr. 1 : 548 ( 1 824) [ ' sphaerocarpon' ] ;Torrey &
Gray, Fl. N. Amer. 1: 163 (1838) ['sphaervcarpon']', A. Gray,
Man. Bot. n. U.S. : 53 ( 1 848); S. Watson, Bibliogr. index N. Amer.
hot. 1 : 1 29 ( 1 878); Coulter inBot. Gaz. 11: 87 ( 1 886); R. Keller in
Engl.&Prantl,Mtf./yZanzen/am.2nded.21: 181 (1925); Small,
Man. s.e.fl.: 871 (1933); Svenson in Rhodora 42: 17 (1940);
Gleason, New Britton & Brown III. Fl. 2: 540 ( 1 952); W.P. Adams
in Contr. Gray Herb. Harv. no. 189: 39 (1962), in J. Elisha
Mitchell scient. Soc. 89: 68 (1973); Steyermark, Fl. Missouri:
1 064 ( 1 970); Utech & Iltis in Trans Wis. Acad. Sci. Arts Lett. 58:
363, map 3 (1970); Mohlenbr., ///. Fl. Illinois fl. pis Hollies to
Loasas: 42 (1978); Cooperrider in Castanea 54: 7, f. 1 (1989).
Type: U.S.A., Kentucky, 'Hab. in Kentucky. Route de Louisville',
n.d. (fl), Michaux s.n. (P-holotype, A-photograph, BM!-micro-
fiche; GH-sketch).

Map 11.

H. nudiflorum sensu Rchb., Icon. Bot. Exot.: 60, t. 87 (1827).
Brathydium sphaerocarpum (Michx.) Spach, Hist. nat. veg. Phan. 5:

444 (1836); K. Koch, Hon. dendrol.: 67 (1853).
B. chamaenerium Spach, Hist. nat. veg. Phan. 5: 445 (1836), in

Annls Sci. nat. (Bot.) II, 5: 365 ( 1 836). Type: U.S.A., Ohio, Herb.

Moser (P-holotype); in ditione 'Miami' civitatis Ohio, 1835 (fl),

Frank s.n. (BM!, K!).

B. chamaerinum Steudel, Nomencl. 2nd ed. 1: 224 (1840), sphalm.
Hypericum chamaenerium (Spach) Steud., Nomencl. 2nd ed. 1: 787

(1840).
H. cistifolium sensu Coulter in A. Gray, Syn. fl. N. Amer. 1: 287

(1897); Small, Man. s.e.fl.: 871 (1933); pro parte uterque excl.

typum.
H. sphaerocarpum Michx. var. sphaerocarpum Svenson in Rhodora

42: 17 (1940), autonym; J.P. Gillespie in Castanea 24: 29 (1958);

Mohlenbr., ///. Fl. Illinois fl. pis Hollies to Loasas: 43, f. 18

(1978).
H. turgidum Small, Fl. s.e. U.S.: 788 (1903), Man. s.e.fl.: 871

(1933); Harper in Geol. Surv. Ala. Monogr. 9: 273 (1928); Rehd.,

Man. cult, trees 2nd ed.: 640 (1940). Type: Alabama, Madison

Co., between Huntsville and Summerville, 7 October 1897 (fr),

Canby 14 (NY-holotype; MO-isotype).
H. sphaerocarpum var. turgidum (Small) Svenson inRhodora 42: 1 7

(1940); J.P. Gillespie in Castanea 24: 29 (1958); Mohlenbr. &

Evans InRhodora 74: 1 46 ( 1 972); Mohlenbr., ///. Fl. Illinois fl. pis

Hollies to Loasas: 43, f. 19 (1978).

Icones: Rchb., Icon. Bot. Exot.: t. 87 (1827); Mohlenbr., ///. Fl.
Illinois fl. pis Hollies to Loasas: 44-^5, ff. 18, 19 (1978).

STUDIES IN HYPER1CUM

Subshrub (stems woody at base) or rhizomatous perennial herb,
0.22-0.58 m tall, erect or decumbent, unbranched or with spreading
to ascending branches from lower half of stem upwards. Stem red-
brown, 2-4-lined, the subfoliar lines more prominent, not becoming
terete; cortex exfoliating in strips. Leaves sessile, 30-70 x 3-1 5 mm,
narrowly elliptic or narrowly oblong to linear, with margin plane to
revolute, slightly to densely glaucous beneath, thinly coriaceous,
persistent, apex obtuse (or sometimes subacute) to rounded, base
narrowly cuneate to parallel; venation: 1 pair of ascending near-
basal laterals andc. 4 pairs of obscure laterals or apparently 1 -nerved,
only midrib prominent; laminar glands dense, small. Inflorescence a
c. 7-70-flowered regular dichasium without accessory flowers,
sometimes with small dichasia from 1-2 nodes below and short
flowering branches from up to a further 6 nodes, the main flores-
cence rounded-corymbiform; pedicels absent or up to 1 .5 mm long;
bracts 2.5-5 mm long, triangular-lanceolate. Flowers (10-)12-15
mm in diam., buds broadly ovoid, apiculate. Sepals 5, 2.5-5 x 1.5-
3 mm, persistent, somewhat unequal, broadly ovate to oblong-elliptic,
obtuse to acute, margin often subrecurved, basal veins 3, unbranched.
Petals 5, bright? yellow, 5-9 x 3-6 mm, 1 .7-2 x sepals, oblanceolate-
elliptic to elliptic, with apiculus lateral, apiculate. Stamens 45-85,
longest 4.5-6.5 mm, 0.65-0.8 x petals, at least some persistent until
fruit matures. Ovary 3-merous, 1 .5-2 x 1-1 .5 mm, ± broadly ovoid,
not lobed, obtuse to acute, 1 -locular with intruding placentae; styles
3, c. 4 mm, 2-2.5 x ovary, remaining appressed in fruit. Capsule 4.5-
8 x 4—7 mm, broadly ovoid to depressed-globose, subapiculate to
rounded. Seeds blackish brown, 2-2.7 mm long, carinate; 'coarsely
reticulate'.

Barren embankments, wet or mesic prairies, limestone outcrops,
cedar glades and sandy stream banks; lowland.

U.S.A., central basin from Iowa, Wisconsin and Ohio to Texas,
Mississippi and Alabama; Florida?.

U.S.A. Alabama: Hale Co., c. 2.9 km S. of Greensboro by Ala 69 1 , 29
May 1 972 (fl), Krai 46840 (BM, MO); Madison Co., see type of//, turgidum;
*Morgan Co., Valhermosa Mt., 28 September 1927 (fr), Harper 39 (A, GH,
MO, NY, US). Arkansas: *Logan Co., 1 2 km SE of Paris, Ms 5344 (MICH,
SMU); Washington Co., Fayetteville, 6 July I9\5(f\), Palmer 8\W(K,MO).
Illinois: ? Co., Beardstown, July 1842 (fl), Geyer s.n. (K); *Champaign Co.,
near Urbana, 3 July \944(ft),Jones 16445 (MO);WinnebagoCo., Rosendal
(Rockford), 12 July 1871 (fl), Cervin s.n. (H). Indiana: *Jasper Co., c. 5 km
NW of junction of Roads 16 and 53, 13 July 1940 (fl & fr), Friesner 14557
(GA, MO, NY, UGA); *Know Co., Vincennes, 14 July 1935 (fl), Hermann
6603 (MO). Iowa: *Black Hawk Co., 20 July 1 929 (fl), Burk 578 (MO); Scott
Co., Davenport, Duck Creek, July 1886 (fl), Fawcett s.n. (BM). Kansas:
*Hempstead Co., McNab, 19 September 1919 (fr), Palmer 16337 (MO);
*Osage Co., c. 5 km S. of Lyndon, Horr & Franklin E323 (FLAS, IND,
NCSC, SMU, US); Wilson Co., 1896 (fl & fr), Holler 631 (JE, MO).
Kentucky: ? Co., Falls of Ohio, 1 842 (fl). Short s.n. (BM. K); ? Co., banks of

119

Kentucky R., July (fl). Peter s.n. (K). Mississippi: *Chicksaw Co., Ray 8548
(MISSA). Missouri: Jasper Co., Joplin, 6 July 1957 (fr), Demaree 39344
(BM); *Madison Co., Black Mountain, S W of Fredricktown, 4 October 1969
(fr), D'Arcy 3832 (MO); St. Francis Co., c. 9 km SE of Cadet, 20 July 1941
(fl & fr),Mewr2088 (BM). Ohio: Franklin Co., Columbus, SciotoR., July (fl
& fr), Lesauereux 29 (BM, K); Hamilton Co., Cincinnati, in ditione 'Miami'
civitatis Ohio, 1835 (fl), Frank s.n. (BM, K). Oklahoma: *Garvin Co., near
Brady, 29 July 1 933 (fl & fr), Palmer 42056 (MO). Pennsylvania: * Alleghany
Co., Glenshaw, 25 June 1921, Bright s.n. (CM). Tennessee: Bedford Co., 8
km N. of Shelbyville, 3 July 1958 (fl), Adams 71 (DUKE, F*, FSU*, IA*.
IND*, MO*, TEX*, VPI*); Wilson Co., Lebanon State Park, 1 80 m, 22 June
1962 (fl), Demaree 45722 (BM). Texas: north-eastern Texas,./z<fc Correll &
Johnston (1970: 64). Wisconsin: Grant Co., 6.4 km NW of Cassville, 7
October 1972 (fl & fr), Nee 5366 (G).

Svenson ( 1 940) recognized a narrow-leaved bushy-branched form
of H. sphaerocarpum as a variety, van turgidum (Small) Svenson,
claiming that it has a well defined (southern) geographical distribu-
tion; and both varieties have been recorded fromTennessee (Gillespie,
1958) and Illinois (Mohlenbrock & Evans, 1972). The variation in
H. sphaerocarpum, however, appears to be continuous, thus prevent-
ing the recognition of any varieties.

H. sphaerocarpum is related to both 17. H. nudiflorum and 1 8. //.
cistifolium. It differs from both in being semi-herbaceous and in
having a central rather than an eastern or south-eastern distribution,
from H. nudiflorum in having narrower leaves, persistent sepals,
smaller flowers and the capsule apex subapiculate to rounded, and
from H. cistifolium in having longer leaves, larger flowers, broader
and obtuse to acute sepals and broader capsules with much larger
seeds.

21. Hypericum adpressum W.P.C. Barton, Comp. Fl. Philadelph.
2: 15(1818);Torrey&Gray, Fl. N.Amer. 1: 159 (1838); Coulter
in Bot. Gaz. 11: 86 (1886), in A. Gray, Syn.fl. N. Amer. 1: 287
( 1 897); B.L. Rob. in Rhodora 4: 1 36, ff. 5-9 ( 1 902); R. Keller in
Engl. & Prantl.Mtf. Pflanzenfam. 2nd ed.21: 1 80 ( 1 935); Small,
Man. s.e.fl.: 871 (1933); Svenson in Rhodora 42: 18 (1940);

Map 12 Sect. 20: 21. H. adpressum • specimens, D records (Kentucky
record incompletely localized); 26. H. tetrapetalum • specimens,
O records.

120

N.K.B. ROBSON

Rehd., Man. cult, trees 2nd ed.: 640 (1940); J.P. Gillespie in
Castanea 24: 28 ( 1958); W.P. Adams in Contr. Gray Herb. Har\>.
no. 189: 41 (1962), in J. Elisha Mitchell scient. Sac. 89: 68
(1973); Radford, Ahles & Bell, Man. vase. fl. Carol inas: 715
(1968); Strasbaugh & Core, Ft. W. Virginia 2nd ed.: 638 (1973);
Mohlenbr., ///. Fl. Illinois fl. pis Hollies to Loasas: 33, f. 13
(1978); M.L. & R.G. Brown, Herb, pis Maryland: 644 (1984).
Type: U.S.A., Pennsylvania, Montgomery Co., 'on the lower
edge of Lansdown grounds, close to the Schuylkill, and not far
above Break's island', c. 1814, Barton s.n. (PH-holotype).
Map 12.

H. fastigiatum Elliot, Sketch hot. S. Carolina 2: 31 (1821). Type:
U.S.A., Georgia, Scriven Co., Elliot s.n. (CHARL-holotype).

H. bonaparteaeW.P.C. Barton, Fl. N. Amer. 3: 95, 1. 106 ( 1 823).Type
as for H. adpressum W.P.C. Barton. Under both species, Barton
cites 'Hypericun No. 6. Bart. Fl. Philad.: 74 (1815)'.

H. adpressum \ar.fastigiatum (Elliot) Torrey & Gray, Fl. N. Amer. 1:
673(1840).

Mvriandra adpressa (W.P.C. Barton) K. Koch, Hon. dendrol.: 66
"(1853).

Brathydium fastigiatum (Elliot) K. Koch, Hort. dendrol.: 66 (1853).

Hvpericum adpressum var. spongiosum B.L. Rob. in Rhodora 4:
136, t. 37 ff. 10-11 (1902); Svenson in Rhodora 42: 19 (1940).
Type: U.S.A., Massachusetts, Bourne, Flax Pond, 15 September
1901 (fr), Kennedy, Williams & Fernald in PI. Exs. Gray. 234
(GH!-holotype; BKL, BM!, F!, MICH, NCSC, PH, TENN, US!,
Z!-isotypes).

H. adpressum forma spongiosum (B.L. Rob.) Fernald in Rhodora
51: 112 (1949); Fernald in Gray's Man. Bot.: 1012 (1950); J.P.
Gillespie in Castanea 24: 28 (1958).

Icones: B.L. Rob. \nRhodora4: t. 37 ff. 5-1 1 (1902); Mohlenbr., ///.
Fl. Illinois fl. pis Hollies to Loasas: 34, f. 13 (1978).

Perennial herb, sometimes woody or spongy at base, (0.2-)0.4-0.8
m tall, erect from creeping rhizomatous base, unbranched until fruit
matures or with 1(?2) pairs of branches below inflorescence and
often axillary leaf clusters. Stems red-brown, 2-lined and ancipitous
above, terete below. Leaves sessile, (15)30-75(-90) x 2-10 mm,
narrowly oblong or linear to narrowly elliptic or lanceolate, with
margin ± revolute, paler but not glaucous beneath, thinly coriaceous,
persistent, apex acute, base narrowly cuneate to parallel, slightly
decurrent; venation: c. 14 pairs main laterals, with subsidiaries
almost equally strong and dense tertiary reticulation, midrib mark-
edly prominent, main laterals less so; laminar glands dense,
medium-sized. Inflorescence c. 1 3-60-flowered regular dichasia,
without accessory flowers or subsidiary branches, rounded-
corymbiform; pedicels 0.5-1.5 mm long; bracts 2-6 mm long,

linear-lanceolate. Flowers c. 10-15 mm in diam., buds ellipsoid.
Sepals 5, (2-)4-7 x 1-1.5 mm, persistent, subequal, ovate-lanceo-
late, acute, margin recurved, often ± deflexed, basal veins 3,
unbranched. Petals 5, bright? yellow, 6-8 x c. 3 mm, 1 .5-2 x sepals,
obovate-oblanceolate, with apiculus obsolete. Stamens c. 60-80,
longest 6-7 mm, 0.85-0.9 x petals, persistent. Ovary 3-merous, 3-
3.5 x 2-2.5 mm, ovoid, not lobed, acute, 1-locular with slightly
intruding placentae; styles 3, (l-)2.5-3 mm long, c. 0.7-0.85 x
ovary, remaining appressed in fruit. Capsule 3.5-6 x 2-4 mm,
ellipsoid to ovoid-ellipsoid, obtuse to rounded. Seeds blackish brown,
0.6-0.7 mm long, slightly carinate; testa scalariform. 2n = 1 8 (n = 9)
(Hoar & Haertl, 1932 (also 'var. spongiosum'); Bostick, 1965).

Marshes, pond margins, wet ditches, bogs (Carolinas); lowland.

U.S.A., east coastal plain from Massachusetts to Georgia (also
inland Ga.), Tennessee, Kentucky?, Missouri and south of L. Michi-
gan.

U.S.A. Connecticut: *New London Co., shore of Shetucket R., 8 August
1 899, Graves s.n. (GH, NEBC). Delaware: New Castle Co., Wilmington (fl),
Tokall s.n. (K); Kent Co., Felton, Upland Meadows, July, Canby s.n. (K).
Georgia: *Dougherty Co., W. of Pretoria, Thome 5709 (GA, IA). Illinois:
*Will Co., Braidwood, Clute20 (NY). Indiana: *JasperCo., 4 km SE ofTefft,
Deam 45934 (GH, IND). Maryland: *Nantucket Co., Nantucket, Almanac
Pond, 14 August 1883 (fl & fr), Churchill 573 (MO). Massachusetts:
Barnstable Co., Bourne, Flax Pond, 15 September 1901 (fr), Kennedy,
Williams & Fernald in PI. Exs. Gray. 233 (BM, K, MO, Z). Missouri: fide
Adams (1959: map 15). New Jersey: no precise locality or date (fl), Barton
s.n. (BM); *Gloucester Co., Hardingville, Long 47122 (PH). New York:
*Suffolk Co., Bridgehampton, Svenson 5202 (TENN). North Carolina: North-
ampton Co., near Margarettaville, 28 July 1 893 (fl), Heller 1 1 55 (GH*, MO*,
PH*, Z). Pennsylvania: Bucks Co., Bristol, 30 July 1 865 (fl), Parker s.n. (K,
MO); see also type. Rhode Island: Washington Co., South Kingstown, 13
August 1878 (f\),Congdon s.n. (BM). South Carolina: Clarendon Co., Dingle
Pond,c. 3. 2 km S. of US 301, c. 3.6 km N. of Marion, 16 September 1967 (fr),
Radford, Bozeman & Leonard 1 1457 (BM, H); *Jasper Co., c. 2.1 km S. of
Tillman, Ahles 15675 (NCU). Tennessee: *Coffee Co., S. of Manchester,
Svenson 8783 (DUKE, MO, PH, WIS). Virginia: *Sussex Co., Stony Creek,
Fernald & Long 10727 (BM, DUKE, GH, MO, US).WestVirginia: Greenbriar
Co., White Sulphur Springs, fide Millspaugh (1892: 338).

H. adpressum in some respects is intermediate in form between 20.
H. sphaerocarpum and 22. H. ellipticum, and its distribution over-
laps those of both species. It is more herbaceous than H.
sphaerocarpum, with rhizomes and shorter fleshier stems that are
sometimes spongy (aerochymatous) at the base, acute leaves and
smaller seeds. For differences between H. adpressum and H.
ellipticum see p. 122.

The variation in H. adpressum is continuous and so there are no
grounds for recognizing the wet-habitat form with fleshy
aerenchymatous stems ('f. spongiosum') as distinct (see Adams,
1962).

22. Hypericum ellipticum Hook., Fl. bor.-amer. 1: 110 (1831);
Torrey & Gray, Fl. N. Amer. 1: 164 (1838); S. Watson, Bibliogr.
index N. Amer. hot. 1: 126 (1878); Coulter in Bot. Gaz. 11: 88
(1886), in A. Gray, Syn. pi. N. Amer. 1: 287 (1897); R. Keller in
Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925);
Fernald, Gray's Man. Bot.: 1012 (1950); Gleason, New Britton
& Brown III. Fl. 2: 539 ( 1952); W.P. Adams in Contr. Gray Herb.
Harv. no. 189: 42 (1962), in/ Elisha Mitchell scient. Soc. 89: 68
(1973); Utech & Iltis in Trans. Wis.Acad. Sci.Arts Lett. 58: 336,
map 4, f. 1 (1970); Scoggan in Publs Bot. natn. Mus. nat. Sci.
Can. no.7(3): 1096 (1979); Mohlenbr., ///. Fl. Illinois fl. pis
Hollies to Loasas: 35, f. 14 (1978); Gillett & Robson in Publs
Bot. natn. Mus. nat. Sci. Can. no. 1 1: 4, tt. 1, 15 ff. 1-2, map 1

STUDIES IN HYPER1CLJM

121

Map 13 Sect. 20: 22. H. ellipticum • specimens, O records.

( 1 98 1 ); Cooperrider in Castanea 54: 7, f. 1 ( 1 989); N. Robson in
Cullen et al., Eur. Gdn Fl. 4: 69 (1995). Type: Canada, 'Canada
to Lake Winnipeg', n.d. (fl), Cleghorn s.n. (K-lectotype, Adams,
1962); Saskatchewan (lat. 54°), 1819-1822 (fl), Richardson s.n.
(K!-syntype, BM!); Ontario, Lake Huron, Pentanguishene, 'ex-
actly on the borders of the United States', Todds.n. (K?-syntype).
The Todd syntype has not been found at K.
Map 13.

H. sphaerocarpum sensuW.P.C. Barton, Comp.fl. Philadelph. 2: 14
(1818), nonMichx. ( 1803).

Brathydium canadense Spach, Hist. nat. veg. Phan. 5: 446 ( 1 836), in
Annls Sci. nat. (Bot.) II, 5: 365 (1836). Type: Canada, Michaux
s.n. (P-syntype); Canada, Lady Dalhousie s.n. (Fl-syntype).

Hypericum brathydium Steud., Nomencl. hot. 2nd ed. 1: 786 ( 1 840).
Type as for Brathydium canadense Spach.

?//. canadense var. oviforme R. Keller in Bull. Herb. Boissier II, 8:
189 (1908). Type: U.S.A., Pennsylvania (not found).

H. ellipticum forma submersum Fassett in Rhodora 41: 376 (1939).
Type: Canada, Ontario, Walford, Pipe Lake, Jack Wilson's Resort,
5 August 1936, Fassett 19172 (WIS-holotype).

H. ellipticum forma foliosum Viet, in Nat. canad. 71: 201 (1944);
Scoggan in Publs Bot. natn. Mus. nat. Sci. Can. no. 7(3): 1096
(1978). Type: Canada, Quebec, Comte de Portneuf, Riviere du
Cap Rouge, 7 September 1941 (st), Marie-Victorin et al. 56602
(MT-holotype; F!, GH!, H!, MO, PH, US!-isotypes).

Icones: Mohlenbr., ///. Fl. Illinois fl. pis Hollies to Loasas: 36, f. 14
(1978); Gillett & Robson in Publs Bot. natn. Mus. nat. Sci. Can., no.
11:6, t. 1 (1981).

Perennial herb somewhat woody at base, 0. 1 1-0.3(-0.5) m tall,
erect from creeping rhizomatous base, unbranched or occasionally
branched below. Stem red-brown, 4-lined, subancipitous above.
Leaves sessile, 1 1-35 x 3-13 mm, rather broadly to narrowly elliptic
or oblong-elliptic or oblanceolate, margin plane to subrevolute,

paler but not always glaucous beneath, lower ones eventually decidu-
ous above base, apex rounded, base cuneate to shallowly
cordate-amplexicaul; venation: 5-7 pairs of main laterals with sec-
ondaries almost equally strong and rather dense tertiary reticulation,
only midrib prominent; laminar glands rather dense, obscure in direct
light, marginal glands dense. Inflorescence ( l-)3-l 5-flowered, regu-
larly dichasial, sometimes with flowering branches from 1-2 nodes
below, corymbiform; pedicels 1-2 mm long; bracts 4-6(-9) mm,
narrowly oblong to linear-lanceolate. Flowers 12-15 mm in diam.,
buds ellipsoid, rounded. Sepals 5(4), 6-7 x 2-3 mm, persistent,
somewhat unequal, oblanceolate, obtuse to rounded, margin plane,
basal veins 3, branched. Petals 5(4), pale? yellow, sometimes tinged
red, 6-8 x 3.5^1 mm, obovate to oblanceolate, apiculus obsolete or
absent. Stamens c. 70-95, longest 4-6 mm, c. 0.7 x petals, persistent.
Ovary 3-merous,(0.8-) 1.5-3. 5 x 1-1. 7 mm, narrowly ovoid, acute, 1-
locular with intruding placentae; styles 3, 1.5-3 mm, 0.8-1 x ovary,
remaining appressed in fruit. Capsule 4—7 x 3.5-5 mm, ellipsoid to
globose, obtuse to rounded. Seeds reddish brown, 0.6-0.7 mm long,
carinate; testa scalariform-reticulate. 2n = 18 (Gillett, 1975, 'c. 18';n
= 9, Hoar&Haertl, 1932), 1 6 ( Love & Love, 1982).

122

N.K.B. ROBSON

Stream, lake and pond margins, river flats, wet meadows and
swamps; lowland to c. 300 m.

Canada (south-western Ontario to southern Newfoundland, extinct
further west? - see Richardson's isotype from 'Saskatchewan'),
U.S.A. (Minnesota to Maine and south to extreme north-eastern
Tennessee, West Virginia, Pennsylvania and northern Delaware).

CANADA. New Brunswick: * Aguance, 1 9 July 1 90 1 (fl & fr), Church-
ill s.n. (MO); *Bass River, 29 July 1 873 (fl), Fowler s.n. (MO). Newfoundland:
Waterford R. between Waterford Bridge and St. John's, 2 August 1911 (fl),
Fernald & Wiegand 5843 (BM, GH*, K, NY*, PH*, US*); St. John's, St.
George's to Port au Port, Codroy R., South Branch, 1-4, July 1949 (fl),
Toumikoski 116 (H). Nova Scotia: Halifax Co., Musquodoboit Harbour,
Petpeswick, 31 July-3 August 1930 (fr), Rousseau 35284 (H, K); Yarmouth
Co., Arcadia, Trefrey's Lake, 29 July 1920 (fl), Fernald & Long 21855 (K).
Ontario: Algoma Distr., Firesand Creek at Hwy 101, 12.8 km E. of Wawa, 7
August 1971, Carton 14702 (H); Kenora Distr., English R., 12.4 km W. of
Trans-Canada Hwy, near Cloven Lake, 7 August 1961 (fl & fr), Baldwin 9378
(H); Thunder Bay Distr., 9.6 km S. of MacDairmid, 0.8 km W. of south end
of Orient Bay, 18 June 1960 (fr), Carton 8313 (BM). Quebec: Cte de
Argenteuil, St-Adolphe d'Howard, 8 August 1 966 (fl & fr), Rolland-Germain
374 17 (BM, H); Chicoutimi Co., vallee de la riviere Ste-Marguerite, 6 August
1964 (fl), Cayouette 7044 (H); Tamiskaming Co., Ottawa R., NE bay of
Grand Lake Victoria, road to L. Granet, 1 5 July 1954 (fl), Baldwin 5870 (K).

U.S.A. Connecticut: *Hartford Co., Southington, 11 July 1897 (fl),
Bissell 344? (MO); *New London Co., Franklin Co., 1 1 July 1906, Wood-
ward s.n. (GH). Illinois: Fulton Co., Lewistown, June 1888 (fl), Repoon s.n.
(BM). Maine: Penobscot Co., Orono, 1 August 1908 (fl), Fernald 235 (BM,
K, MO*, Z). Maryland: *Garrett Co., vicinity of Mount Lake Park, Steele 56
(GH, US). Massachusetts: Essex Co., Florence (Northampton), 24 July 1977
(fr), Ahles 8429 1 (BM, H): Hampshire Co., Andover, July 1 882 (fl), Blake s.n.
(H). Michigan: *Chippewa Co., near Saulte Sainte Marie, 14 August 1910(fl
& fr), Churchill s.n. (MO); *Chippewa Co., Sugar I., McVaugh 875 1 (MICH,
NCSC). Minnesota: *Lake Co., along Baptism R., N. shore of L. Superior,
Moore & Huff 18742 (GH, IND). New Hampshire: Hillsboro Co., Pelham,
Long Pond, 14 July 1927 (fl), Beanie s.n. (BM, K); Cheshire Co., Surry,
Ashuelot R., 9 August 1972 (fr), Boufford 7570 (MO*, Z). New Jersey:
Madison Co., Lennox, Oneida Lake, 24 July 1901 (f\),Haberer 147(K).New
York: *Hamilton Co., Coles Landing, 6 September 1926 (fr), Wiegand 16674
(GH, MICH, MO); no precise locality or date (fr), Gray s.n. (BM, K). Ohio:
fide Adams (1959: map 12) and Cooperrider (1989: f. 1). Pennsylvania:
Fayette Co., Ohio Pyle, 29 June 1 902 (fl), Shafer 277 (BM); Forest Co., c. 3.2
km S. of Duhring along Spring Creek, 26 August 1955, Henry s.n. (CM).
Rhode Island: Providence Co., Providence, no date (fl), Olney s.n. (K).
Tennessee: Johnson Co., Shady Valley Bog (fide Gillespie, 1958); *10 km
SW of Mountain City, Shanks & Sharp 7090 (TENN). Vermont: *Franklin
Co., Binghamville, Fairfax, 20 September 1964 (fr), Seymour & Charette
22471 (MO); Windham Co., Jamaica, 13 July 1937 (fl), H. & E. Moldenke
9914 (BM, *MO). West Virginia: *Upshur Co., 6 July 1 895 (fr), Pollock s.n.
(MO). Wisconsin: Florence Co., BruleR., 1 July 1964 (f\),Iltis 22194 (BM);
*SawyerCo., near Hay ward, 1 1 September 1925 (fr), Palmer 28628 (MO).

H. ellipticum is related to H. sphaerocarpum but differs from it inter
alia by the shorter herbaceous rhizomatous habit, shorter leaves and
smaller seeds. A submerged aquatic form with elongate simple
stems and shorter, ovate to orbicular leaves (f. submersum Fassett)
and one with the axillary branches grown out after fertilization (f.
foliosum Marie- Victorin) seem scarcely worth recognizing.

Subsect. 4. Brathydium (Spach) R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 3(6): 214 (1893)
[ 'Eubrathydium '].

Brathydium Spach, Hist. nat. veg. Phan. 5: 442 ( 1 836), in Annls Sci.
nat. (Hot.) II, 5: 365 (1836). Type: see below.

Type: B. grandiflorum Spach, nom. illegit. =Hypericumdolabriforme
Vent, (lectotype, Y. Kimura, 1951).

Shrubs or subshrubs with leaves either articulated at base and

deciduous (Sp. 23) or not articulated and persistent or deciduous
above base (Spp. 23, 24); inflorescence-branching dichasial, mainly
acropetal; sepals 5, very unequal to subequal, persistent; petals 5;
stamens 120-200, deciduous (Sp. 23) or persistent (Sp. 24); styles
and placentae 3(4), placentation incompletely axile (Sp. 23) or
parietal (Sp. 24). Species 23-24.

Kimura (1951) selected his own combination, Brathydium
dolahriforme (Spach) Y. Kimura, as lectotype; but Spach's name,
even though illegitimate, should have been used for the species
selected.

23. HypericummyrtifoliumLam.,£/icvc/. 4: 180(1797);Choisyin
DC., Prodr. 1: 547 (1824); Torrey & Gray, Fl. N. Amer. 1: 161
(1 838); Chapm., Fl. South. U.S.: 40 ( 1 865); S. Watson, Bibliogr.
index N. Amer. hot. 1: 127 (1878); Coulter in Bot. Gaz. 11: 85
(1886), in A. Gray, Syn.fl. N. Amer. 1: 286 (1897); R. Keller in
Engl. & Prantl, Nat. Pflanzenfam. 2nded.21: 180(1925); Small,
Man. s.e.fl.: 872(1933); Svenson \nRhodora42: 19 (1940);W.P.
Adams inContr. Gray Herb. Harv. no. 189: 35 (1962), in/ Elisha
Mitchell scient. Soc. 89: 68 (1973); R.C. Clark in Ann. Mo. hot.
Gdn 58: 209 (1971); R. Long & Lakela, FL Trop. Florida: 607
(1971); Godfrey & Wooten, Aquatic & wetland pis s.e. U.S.
Dicots: 341, f. 154 (1981); Clewell, Guide vase, pis Florida
Panhandle: 372 (1985); Godfrey, Trees, shrubs & woody vines n.
Florida, etc.: 356, f. 169 (1988). Type: U.S.A., without precise
locality, Herb. B. de Jussieu (P-LA-holotype; GH-photograph).

Map 14.

H. glaucum Michx., Fl. bor-amer. 2: 78 (1803); Pursh, FL Amer.

sept. 2: 376 (1814); Elliott, Sketch hot. S. Carolina 2: 32 (1821);

Choisy, Prodr. monogr. Hyperic. : 46 ( 1 821 ), in DC., Prodr. 1: 547

(1824). Type: U.S.A., Florida, n.d. (fl & fr), Michaux s.n. (P-

holotype, BM!-microfiche).
H. rosmarinifolium sensu Choisy, Prodr. monogr. Hyperic.: 45

(1821), in DC., Prodr. 1: 547 (1824).
H. sessiliflorum Willd. ex Spreng., Syst. Veg. 3: 346 (1826). Type;

U.S.A., 'inAmericaborealis', ?(B-WILLD-holotype,BM!-micro-

fiche).
Myriandra glauca (Michx.) Spach, Hist. nat. veg. Phan. 5: 442

(1836), in Annls Sci. nat. (Bot.) II, 5: 365 (1836).
Brathydium myrtifolium (Lam.) K. Koch, Hon. dendrol.: 67 (1853).

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 356, f.
169(1988).

Shrub (or subshrub?), 0.3-1 m tall, mostly 1 -stemmed, erect from
woody caudex, often unbranched below inflorescence, sometimes
with branches in upper half, ascending. Stems glaucous green, 4-

STUDIES IN HYPERICUM

123

Map 14 Sect. 20: 23. H. myrtifolium • specimens, O records; 24. H.
dolabriforme • specimens, D records.

lined and ancipitous when young, soon 4-lined and rounded, becom-
ing reddish brown and 2-lined to terete; cortex exfoliating in strips;
bark greyish, becoming corky, thick Leaves sessile, spreading,
evergreen, (8-) 13-40 x (5-)7-20 mm, oblong-ovate to triangular-
lanceolate, margin recurved (especially when dry), paler and usually
± glaucous beneath, sometimes also glaucous above, coriaceous,
eventually deciduous at or near semi-articulated base, apex rounded
to obtuse or sometimes acute, base subcordate to cordate, ±
amplexicaul; venation: 3-4 main laterals, with laxly reticulate sec-
ondaries, tertiaries not visible; laminar glands very dense.
Inflorescence 7-c. 30-flowered, regularly dichasial, widely branched,
sometimes with 1-3 accessory flowers at apical node, with solitary
flowers or 3-7-flowered dichasia or flowering branches up to 3
nodes below, the whole hemispherical to subcorymbiform; pedicels
to 3 mm long or absent; bracts foliar, reduced. Flowers (15-)20-25
mm in diam.; buds ovoid. Sepals 5, 5-8 x 2-4.5 mm, enlarging
somewhat in fruit, imbricate, unequal to subequal, ovate to lanceo-
late, becoming foliaceous, acute, margin recurved; basal veins (3)5,
branching and reticulating distally. Petals 5, bright yellow, becom-
ing apically recurved, 8-15 x 4.5-6 mm, 1 .5-2 x sepals, obovate to
oblong-oblanceolate, with apiculus lateral, obtuse. Stamens c. 200,
longest 5-9 mm, c. 0.6 x petals, deciduous. Ovary 3(4)-merous, 3-
4 x 1.3-3 mm, narrowly pyramidal-ovoid, acute, placentation
incompletely axile; styles 3(4), 4-5 mm, 1 .2-1 .3 x ovary, separating
above in fruit. Capsule 5-6 x 3-4 mm, pyramidal-ovoid, 3(4)-lobed
or 3(4)-gonous. Seeds blackish brown, c. 1 mm long, narrowly
carinate; testa shallowly linear-reticulate. 2n = 18 (n = 9, Adams in
Robson & Adams, 1968).

Moist pine flatwoods, grass/sedge bogs, margins of evanescent
ponds and low roadside ditches, on sandy or peaty soil; lowland.

U.S.A. (coastal plain from South Carolina? or Georgia to south-
eastern Mississippi, including most of peninsular Florida).

U.S.A. Alabama: Baldwin Co., just N. of junction 1-10 by AL 59, 9.6
km S. Stapleton, 8 June 1971 (fl & e. fr), Krai 43081 (BM): Mobile Co.,

Range Line readjust N. of AL 163, S. of Mobile, 26 May 1973 (fl), Boufford
& Ahles 9423 (BM, MO*). Florida: Duval Co., near Jacksonville, June 1 894?
(fl & fr), Curtiss 265 (BKL*, BM, F*, FR, G, GH*, K, MISSA*, NY*, PH*,
US*); Lake Co., near Eustis, 1-15 May 1894,Mw/i708(G, K);Wakulla Co.,
US 98c. 20.8 kmW. of Newport, 27 June 1 959 (fl), Dress & Readll53 (BM).
Georgia: Sumter Co., 31 August 1900 (fr). Harper 548 (BM, K, MO*);
Thomas Co., 4.8 km N. of Pavo, 13 June 1 968 (fl), Adams 28 (K). Mississippi:
*Jackson Co., Belle Fontaine Point, Diener 915 (MISSA); *Jackson Co.,
near Ocean Springs, May 1859 (fl), Hilgard s.n. (MO).

Small (1933) recorded H. myrtifolium from South Carolina, and
there is a specimen in Herb. De Candolle (G-DC) labelled 'Carol,
merid., Fraser'; but this species is not treated by Radford, Ahles &
Bell (1968).

Adams ( 1 962) included H. myrtifolium in subsect. Centrosperma,
and the leaves do have a groove at the base of the midrib beneath. It
does not extend along the rest of the lamina base, however, so that the
leaves sometimes leave a small zone behind when they fall. In
addition, the sepals are persistent after fruit dehiscence, but the
stamens are deciduous with the petals and the placentation is incom-
pletely axile. All these characters suggest that H. myrtifolium could
be intermediate between subsections Centrosperma and
Suturosperma, but the habit, leaf shape, larger sepals and numerous
stamens indicate a direct relationship with 1. H.frondosum.

24. Hypericum dolabriforme Vent., Descr. pi. nouv.: 45 & f.
( 1 800); Pursh, FI.Amer. sept. : 378 ( 1 8 1 4); Choisy, Prodr. monogr.
Hyperic.: 45 (1821), in DC., Prodr. 1: 547 (1824); Torrey &
Gray, Fl. N. Amer. 1: 162 (1838); Steud., Nomencl. hot. 2nd ed.
1: 787 (1840); S. Watson, Bibliogr. index N. Amer. hot. 1: 126
(1 878); Coulter in Bot. Gaz. 11: 87 ( 1 886), in A. Gray. Syn.fl. N.
Amer. 1: 287 ( 1 897); Britton & Brown, lll.fl. n. states 2nd ed. 2:
532, f. 2889 ( 19 1 3); R. Keller in Engl. & Prantl, Nat. Pflanzenfam.
2nd ed. 21 : 1 8 1 ( 1 925); Small, Man. s.e.fl. : 87 1 ( 1 933); Svenson
in Rhodora 42: 16 (1940); J.R Gillespie in Castanea 24: 29
(1958); W.P. Adams in Contr. Gray Herb. Harv. no. 189: 40
(1962), in J. Elisha Mitchell sclent. Soc. 89: 68 (1973). Type:
U.S.A., Kentucky, 'sur les collines tres arides de Kentucky', n.d.
(fl), Michaux (G!-holotype).

Map 14.

H. procumbens Desf. ex Willd., Sp. pi. 3: 1450 (1802). Type:
U.S.A., 'Amer. Sept.', ? in Herb. Willd. 14424 (B-WILLD-
holotype, microfiche!).

H. procumbens Michx., Fl. bor.-amer. 2:81 (1 803); Pursh, Fl. Amer.
sept.: 379 (1814); Choisy, Prodr. monogr. Hyperic.: 45 (1821), in
DC., Prodr. 1: 547 (1824). Type: U.S.A., Kentucky, 'in aridis
collibus Kentucky', n.d. (fl), Michaux s.n. (P-holotype, BM!-
microfiche). Perhaps this is from the same collection as the type
of H. dolabriforme Vent.

Brathydium grandiflorum Spach, Hist. not. veg. Phan. 5: 443 ( 1 836),
in Annls Sci. not. (Bot.) II, 5: 365 ( 1 836); K. Koch, Hort. dendrol:
66 (1853), nom. illegit. (Art. 63). Type as for Hypericum dola-
briforme Vent.

Streptalon dolabriforme (Vent.) Raf., Fl. Tellur. 3: 80 (1837).

Hypericum bissellii B.L. Rob. in Rhodora 4: 136, t. 37 ff. 1-4
(1902). Type: U.S.A., Connecticut, Southington, 30 July 1901
(fl), Bissell 4025 (GH!-holotype), see p. 124

Brathydium dolabriforme (Vent) Y. Kimura in Nakai & Honda,
Nova fl. jap. 10:24(1951).

Icon: Vent., Descr. pi. nouv.: 45 & f. (1800).

Subshrub 0. 1 5-0.5 m tall, decumbent and woody (but not rooting) at
base, with short or ± elongate branches at base or throughout stem.

124

N.K.B. ROBSON

Stem green?, 4-lined and ancipitous above, 2-lined to terete below;
cortex exfoliating in strips. Leaves sessile, widely spreading: lamina
(main stem) 20-35 x 3-5 mm, linear-elliptic or linear-oblong to
linear, margin recurved to revolute, pale or slightly glaucous? be-
neath, subcoriaceous, the lower deciduous slightly above base, apex
obtuse to acute, base narrowly cuneate to rounded; venation: only
midrib apparent; laminar glands dense. Inflorescence (l-)3-c. 20-
flowered, regularly dichasial, ± widely branched, without accessory
flowers, rarely with single flowers from axil below, the whole
obconic; pedicels 1.5-2 mm long; bracts foliar, reduced, oblong to
lanceolate. Flowers c. 15-20 mm in diam.; buds ± ellipsoid, acumi-
nate. Sepals 5, 5-8(-15) x 2-3(-8) mm, not enlarging in fruit?,
imbricate, very unequal, ovate-lanceolate to lanceolate, ± folia-
ceous, acuminate, margin distally revolute; basal veins 3, laterals
sometimes branched. Petals 5, 'golden' yellow, 10-13 x 4-5 mm, c.
1.6-2 x sepals, curved-dolabriform, with apiculus conspicuous,
termino-lateral, acute. Stamens 120-200, longest 5-7 mm, c. 0.5 x
petals. Ovary 3-merous, 2.5-3 x 1 .5-2 mm, ovoid-conic, acuminate,
placentation parietal; styles 3, 3.5-4 mm, 1.35-1.4 x ovary, some-
times separating above in fruit. Capsule 4— 7(-9) x 3-3. 5(-4) mm,
ovoid-conic, rostrate, 3-gonous above. Seeds reddish, 1.5-1.8 mm
long, carinate; testa reticulate-scalariform. 2n = 1 8 (n = 9, Adams in
Robson & Adams, 1968).

Limestone outcrops, cedar glades and dry rocky stream-beds; low-
land to c. 500 m in Georgia.

U.S.A. (from extreme southern Indiana and north-central Kentucky
southward through eastern Tennessee to north-western Georgia;
probably introduced into Connecticut).

U.S.A. Connecticut: see H. bissellii above, probably introduced. Geor-
gia: ? Co., east base of Pigeon Mts, 300 m, 3 August 1900 (fl & fr), Harper
359 (BM, K); *Catoosa Co., 16 km W. of Ringgold, Cronquist 5614 (GA,
GH, IND, MO, NY, PH, SMU, UCA, US); Walker Co., Chickamauga, 1 1 July
1899 (fl). Herb. Biltmore 6474a (JE, Z). Indiana: *Crawford Co., vicinity of
Wyandotte Cave, 1 3 July 1 899, Blatchley s.n. (IND). Kentucky: * Nelson Co.,
19.2 km S. of Bardstown, McFarland 50 (DUKE, GH, IND, MO, NY, PH,
TENN, WIS); * Warren/Simpson Co. line, on US 3 1 W, 30 June 1969 (fl & fr),
Conrad 236 (MO); *Wayne Co., Monticello, Smith & Hodgdon 401 3 (F, GH,
NY, US); ? Co., Estival, 1 83 1 (fl), Rafinesque s.n. (G).Tennessee: Meigs Co.,
nearDecatur, 13 July 1934(fl),S/wr;;<& Underwood 2293 (BM); Knox Co.,
near Mascot, 30 June 1958 (fl & fr), Adams 61 (K, MO*).

H. dolabriforme is a relict species of which the affinities are not
immediately apparent. Adams (1962: 41) claimed it to be most
closely related to 20. H. sphaerocarpum, which is similarly semi-
woody with linear leaves, axillary leaf-clusters, an almost wholly
terminal dichasial inflorescence and somewhat unequal sepals. The
capsules of//, dolabriforme, however, are much bigger, the inflores-

cence branches more widely spreading, the sepals much more
unequal and recurved-acuminate (rarely subrecurved and obtuse to
acute), the petals curved-dolabriform, the stamens more numerous
(120-200, not 70-95) and the seeds much larger. All these charac-
ters, except the linear leaves and large seeds, can however be derived
easily from those of 23. H. myrtifolium; and the leaves, though
narrow, are of similar texture and colour. The larger seeds need not
be an insuperable obstacle to a relationship with H. myrtifolium,
which seems to be directly related to 1 . //. frondosum like the 4-
petalled 'Ascyrum' species.

I agree with Adams (1962: 41 ) that H. bissellii is a synonym of//.
dolabriforme. Robinson's illustration could be of//, sphaerocarpum,
but the type specimen clearly belongs to H. dolabriforme. Consider-
ing that this species is otherwise confined to west or south of the
Appalachians, the record from a street in a Connecticut town is
likely to have resulted from an introduction.

Subsect. 5. Ascyrum (L.) N. Robson, stat. nov.

Ascyrum L., Sp. pi. : 787 (1753), Gen. pi 5th ed.: 342 ( 1 754) excl. A.
villosum L. et A. crux-andreae L. pro parte quoad syn.; Engl. in
Engl. & Prantl, Nat. Pflanzenfam. 3(6): 208 (1893), op. cit., 2nd
ed. 21: 174 (1925) pro parte, excl. A. filicaule Dyer. Type: A.
hypericoides L. (= Hypericum hypericoides (L.) Cr.) (lectotype -
Britton & Brown, 1913: 528).

Hypericoides Adans., Fam. PL 2: 443, 616 (1763); Lam., Tabl
encycl.: t. 644 (1796). Type: Hypericum hypericoides (L.) Cr. (=
Hypericoides perforata Poir.) (lectotype - N. Robson, 1977a).

Ascyrum a. Ascyrum (L.) Endl., Gen. PL: 1032 (1840), autonym
status ignot.

Type: H. hypericoides (L.) Cr. (see above).

Shrubs or wiry shrublets with leaves articulated at base (Spp. 25, 26)
or not, deciduous sooner or later at or above base; inflorescence-
branching dichasial and/or pseudo-dichotomous, basipetal and/or
acropetal; sepals 4, markedly unequal, persistent; petals 4; stamens
30-100, persistent; styles and placentae 2-4, placentation parietal.
Species 25-29.

25. Hypericum crux-andreae (L.) Cr., Inst. rei herb. 2: 520 ( 1 766);
N. Robson inTaxon 29: 272 (1980); Godfrey &Wooten, Aquatic
& wetlandpls s.e. U.S., Dicots: 340 ( 1981 ); Clewell, Guide vase,
pis Florida Panhandle: 371 (1985); Godfrey, Trees, shrubs &
woody vines n. Florida, etc.: 350, f. 165 (1988); N. Robson in
Cullen et al., Eur. Gdn Fl. 4: 69 (1995). Type: U.S.A., Virginia,
Clayton 230 (LINN 944/1 -lectotype - N. Robson, 1980).

Fig. 18A, Map 15.

Ascyrum crux-andreae L., Sp. pi.: 787 (1753) excl. syn., Sp. pi. 2nd
ed.: 1107 (1763) excl. syn. Plukenet.; Lam., Encycl. 1: 285
(1783) pro parte, excl. syn. Plukenet. et Raii.

Hypericum tetrapetalum [var.] p sensu Lam., Encycl. 4: 153 (1797),
fide microfiche P-LA 73/7.

Ascyrum stans Michx. ex Willd., Sp. pi. 3: 1473 (1802); Michx., Fl.
bor.-amer. 2: 77 ( 1 803); Vent., Jard. Malmaison 2: t. 90 ( 1 805) pro
parte, excl. syn. Hypericum tetrapetalum Lam.; Choisy, Prodr.
monogr. Hyperic.: 61 (1821) pro parte, excl. [var.] p, in DC.,
Prodr. 1: 555 (1824) pro parte excl. syn. H. tetrapetalum Lam.;
Spach, Hist. nat. veg. Phan.5: 457 (1836), inAnnlsSci. nat. (Bot.)
II, 5: 368 (1836); Torrey & Gray, Fl. N. Amer. 1: 157 (1838); A.
Gray, Gen. Amer. bor. 1: t. 91 (1848); Chapm., Fl. South. U.S.: 39
( 1 865); Coulter in Bot. Gaz. 11:81(1 886), in A. Gray, Syn. Fl. N.

STUDIES IN HYPER/CUM

125

Map 15 Sect. 20: 25. H. crux-andreae • specimens, O records.

Amer. 1: 283 ( 1 897); R. Keller in Engl. & Prantl, Nat. Pflanzenfam.
2nded. 21: 174(1925); Small, Man. s.e.fl.: 868 (1933); Fernald,
Gray's Man. Bot.: 1007 (1950); W.P. Adams in Rhodora 59: 88,
map 4 (1957); J.P. Gillespie in Castanea 24: 27 (1958); R.A.
Vines, Trees, shrubs & woody vines ofS. W. : 732 ( 1 960); Correll &
Johnston, Man. vase, pis Texas: 1060 (1970); K.G. & M.L.
Brown, Woody pis Maryland: 233 (1972). Type: U.S.A., 'Hab. in
Carolina', Michaux (P-holotype).

A. hypericoides sensu W.T. Aiton, Hort. Kew. 2nd ed.: 430 (1812);
Elliott, Sketch hot. S. Carolina 2: 22 (1821).

A. amplexicaule sensu Pursh, Fl. Amer. sept. 2: 374 (1814) pro parte,
quoad syn. A. stans Willd.

A. grandiflorum Raf., Fl. ludov: 87 (1817). Type: U.S.A., Louisiana
(no specimen found).

Hypericoides crux-andreae (L.) Poir., Tab. Encycl. 3: 201 , t. 644 f. 1
(1823).

Ascvrum stans [var.] P ohovatum Chapm. ex Torrey & Gray, Fl. N.
Amer. 1: 157 (1838); Chapm., Fl. South. U.S.: 39 (1865). Type:
U.S.A., Florida, 'Middle Florida', Chapman s.n. (NY-holotype).
A. simplex Zeyh. ex Turcz. in Bull. Soc. Nat. Moscou 31(1): 389
( 1 858). Type: U.S.A., Pennsylvania, Bethlehem, n.d. (fl.), Zeyher
s.n. (KW-holotype; BM!-photograph).

A. cruciatum St-Lag. in Annls Soc. hot. Lyons 7: 69 (1880), nom.
illegit. (Art. 63). Type as for A. crux-andreae L.

A. cuneifolium Chapm., Fl. South. U.S. 2nd ed., Suppl. 2: 680
(1892), op. cit., 3rd ed.: 56 (1897); Small, Fl. s.e. U.S.: 785
(1903), Man. s.e.fl.: 868 (1933); Merrill in Castanea 13: 66
(1948). Type: Florida, 1835, Chapman (NY!-holotype).

Hypericum stans (Michx.) Adams & Robson in Rhodora 63: 15
(1961); W.P. Adams in Contr. Gray Herb. Harv. no. 189: 44
(1962), in J. Elisha Mitchell sclent. Soc. 89: 68 (1973); Radford,
Ahles & Bell, Man. vase, pis Carolinas: 1 1 1 (1968); R.C. Clark in
Ann. Mo. hot. Gdn 58: 209 (1971); R. Long & Lakela, Fl. imp.
Florida: 607 (1971).

H. peltatum sensu T. & M. Eisner &Aneshansley in Procnatn.Acad
Sci. U.S.A. 70: 1002 (1973), nomen.

Icones: Vent., Jard. Malmaison 2: t. 90 (1805); Godfrey, Trees,
shrubs & woody vines n. Florida, etc.: 350, f. 165 (1988).

Shrub 0.1-1.0(-1.35) m tall, erect to suberect or rarely decumbent
and rooting at base, unbranched or rarely sparsely branched below
inflorescence (at least until fruiting), branches suberect. Stems

becoming red-brown, 2-4-lined and ancipitous when young, soon
narrowly 2-winged; cortex exfoliating in thin strips or flakes; bark
thin, reddish brown, not corky. Leaves sessile, ascending to spread-
ing, (12-)18-30(-36) x (6-)8-12(-16) mm, oblong to elliptic or
rarely obovate to oblanceolate or triangular-ovate, margin plane to
subrecurved, paler beneath, sometimes slightly glaucous on both
sides, coriaceous, eventually deciduous at basal articulation, apex
rounded to obtuse, base rounded to truncate or rarely slightly
cordate-amplexicaul; venation: up to 3 pairs of laterals sometimes
visible; laminar glands dense. Inflorescence l-3(-7)-flowered from
\-4 nodes, sometimes with flowering branches from up to 4 nodes
below, the whole ± narrowly cylindric to narrowly pyramidal or
occasionally with one pair of pseudo-dichotomous branches; pedicels
3-5 mm long; bracts foliar; bracteoles triangular-lanceolate. Flow-
ers 20-30 mm in diam.; buds compressed-subglobose. Sepals 4,
markedly unequal, not? enlarging in fruit; outer 9-1 7(-20) x 9-1 4(-
18) mm, broadly ovate to circular, apiculate to obtuse or rounded,
base cordate, basal veins (3)5-7, midrib often branched; inner 7-14
x 2-3(^4-) mm, narrowly elliptic to lanceolate, acute to subacute,
basal veins 3(-5), midrib sometimes branched. Petals 4, bright
yellow, 1 1-18 x 6-10(-12) mm, c. 05-1.2 x outer sepals, obovate,
with apiculus lateral, acute. Stamens 80- 100, longest 7-8 mm, 0.45-
0.65 x petals. Ovary 3(4)-merous, (3.5-)4-5 x 1 .5-2.5 mm, narrowly
ellipsoid-ovoid, acute, placentation parietal; styles 3(4), (1-) 1.5-2. 5
mm, 0.35-0.5 x ovary, divergent. Capsule 7-9(-10) x 5-6.5 mm,
narrowly ellipsoid-ovoid, obtuse (or apiculate fide Small), scarcely
lobed. Seeds blackish brown, 0.8 mm long, ecarinate; testa shallowly
scalariform. 2n = 18 (n = 9, Adams in Robson & Adams, 1968).

Moist to dry pine savannahs and flatwoods, meadows, bogs, marshes,
ditches, shores of ponds and lakes, on sandy soil; lowland.

U.S.A. (eastern Texas and south-eastern Oklahoma to Florida and
northwest to New York (Long Island), New Jersey and eastern
Pennsylvania).

U.S.A. Alabama: *Barbour Co., by L. Eufaula, 8 km S. of Eufaula, 1 1
September 1968 (fr), Krai 33219 (MO); Talladega Co.?, 'supra Talladega ad
fl. Coosa', Sept. 1843 (fr), Rugel s.n. (BM); *Mobile Co., Graves 697 (MO,
US). Arkansas: *ClarkCo.,Arkadelphia,60m, I October \93%(fT),Demaree
18422 (MO); Hot Springs Co., Malvern, 4 September 1915 (fl), Palmer&463
(K, MO*); no precise locality or date, Leavenworth s.n (K). Delaware: *New
Castle Co., Tatnall 1959 (GH, PENN); Sussex Co., Ellendale, August 1867
(fl), Canby s.n. (H). Florida: Franklin Co., W. of Fla 379, c. 19 km S. of
Liberty-Franklin county line, 9 November 1 963 (fr). Ward & Ford 3645 (BM,
FLAS*): Hillsborough Co., NE Tampa, Industrial Park, W. of 46th St.,
intersected by Linebaugh Ave., 13 November 1963 (fr), Lakela 26666 (BM,
USF* ). Georgia: BullochCo., SE of Statesboro, 4.9 km N. of Georgia 46 on
co. road 204, then right for 1.3 km, c. 65 m, 16 September 1982 (fl & fr),
Boufford, Bartholomew & Spongberg 23137 (A*. BM, MO*): Thomas Co.,
c. 5 km NW of Pavo. 10 August 1958 (fl), Adams 159 (K). Kentucky:

126

N.K.B. ROBSON

Fig. 18 A. H. crux-andreae: (a) habit; (b) leaf (part); (c) outer sepal; (d) inner sepal; (e) petal; (0 capsule. B. H. tetrapetalum: (g) habit. C. H.
edisonianum: (h) habit. D. H. suffruticosum: habit (a, g-i x '/2; b x 2: c-f x 3). A. Boufford2\405. B. Skean 1600. C. Judd 25 17. D. Curtiss 246.

STUDIES IN HYPER1CUM

*McCreary Co., Bauer Road, 24 July 1941 (fl), McFarland & James 48
(DUKE, IND, MO, PENN, TENN, US, WVA). Louisiana: *Beauregard Par.,
Correll&Correll96%\ (DUKE,F,GH); *RapidesPar., l896(fl),Do£fao/is.n.
(MO); St. Tammany Par., Covington, June-August 1 832 (fl), Dmmmond s.n.
(BM, K). Maryland: "Prince George Co., Hyattsville, 14 August 1910 (fl),
Dowell 6464 (GH, MO, NY); *Worcester Co., 0.8 km N. of Ocean City, 1 1
September 1936 (st), Norton s.n. (M). Mississippi: Harrison Co., Gulfport, 5
September 1900 (fl), Tracy 87 (BM); *Ocean Springs, 22 August 1891 (fl),
Seymour 91 822 (MO). New Jersey: Atlantic City, 30 August 1 882 (fl), Parker
s.n. (K); Burlington Co., c. 9.6 km W. of Chatsworth, 20 August 1948 (fl),
Lawrence & Dress 528 (BM). New York: see Adams (1957). North Carolina:
"Macon Co., Highlands, 31 July 1975 (fr), Boufford & Wood 77755 (MO);
Onslow Co., 8.8 km S. of N.C.4 1 on County road 1 003, S. of Comfort and NE
of Richmond City, 22 August 1979 (fl & fr), Boufford 21405 (BM, CM*).
Oklahoma: *Le Flore Co., Palmer 20595 (GH). Pennsylvania: *Bucks Co.,
28 August 1864, Diffenbaugh s.n. (GH); Philadelphia Co., near Philadelphia,
August (fr), Griffith 136 (BM). South Carolina: Marlboro Co. 2.4 km E. of
Wallace, 10 August 1956 (fl), Radford 15625a(K); Oconee Co., Whitewater
R. 3.2 km NW of Jocasee, 2 September 1956 (fl), Radford 17892 (H).
Tennessee: "Coffee Co., Svenson 4246 (GH, IND, PH); *Franklin Co., 10
September 1898 (fl), Eggerts.n. (MO); Grundy Co., 6.4 km N. of Palmer by
Savage Gulf off Collins School Road, 2 September 1971 (fr), Krai 43739
(BM). Texas: "Houston Co., Latexo, 22 September 1917 (fl), Palmer 12819
(MO); *Smith Co., along SabineR., near Gumwood, 27 September 1926 (fr),
Palmer 31727 (MO). Virginia: Norfolk Co., prope Norfolk, August 1840(fl),
Rugel 202 (BM); "Princess Anne Co., near Virginia Beach, Heller 1268 (F,
GH, NY, PENN, PH, US).

H. crux-andreae, with its tetramerous perianth whorls and very
unequal sepals, is directly related to 1. H.frondosum, in which the
sepals are very unequal and both perianth whorls are quite often
tetramerous. It differs from that species in having (as well as the
constantly tetramerous perianth) sessile leaves, a lower habit and
smaller flowers at more stem nodes.

Adams (1957) showed that Chapman's Ascyrum cuneifolium (A.
stans var. obovatum Chapm. ex Torrey & Gray) cannot be separated
from typical//, crux-andreae (=A. stans). The low, several-stemmed
form with cuneate leaves, longer-pedunculate flowers and shorter
sepals varies in the direction of 28. H. suffruticosum, but there is no
overlap in variation in these taxa.

26. Hypericum tetrapetalum Lam., Encycl. 4: 153 (1797); Adams
&Robson \nRhodora63: 15 (1961);W.P. Adams inContr. Gray
Herb. Harv. no. 189: 45 (1962), in/ Elisha Mitchell sclent. Soc.
89: 68 (1973); R. Long & Lakela, Fl. trap. Florida: 606 (1971);
Godfrey & Wooten, Aquatic & wetland pis s.e. U.S. Dicots: 340
( 1 98 1 ); Clewell, Guide vase, pis Florida Panhandle: 372 ( 1 985);
Godfrey, Trees, shrubs & woodv vines n. Florida, etc.: 348, f.
164 (1988). Type: U.S.A., Florida, 'Hab. in Florida', ? (P-LA-
holotype, GH- photograph).

Fig. 18B, Map 12.

Ascyrum amplexicaule Michx., Fl. bor.-amer. 2: 77 (1803); Pursh,
Fl.Amer. sept. 2: 374 ( 1 8 1 4); Elliott, Sketch hot. S. Carolina 2: 23
(1821); Choisy in DC., Prodr. 1: 555 (1824); Spach, Hist. not.
veg. Phan. 5: 457 (1836), in Annls Sci. nat. (Bot.) II, 5: 368
(1836); Torrey & Gray, Fl. N. Amer. 1: 156 (1838); Chapm., Fl.
South. U.S.: 39 (1865); Coulter in Bot. Gaz. 11: 81 (1886), in A.
Gray, Syn. fl. N. Amer. 1: 283 (1897). Type: U.S.A., Florida,
without precise locality, Michaux s.n. (P-holotype:, BM!-micro-
fiche).
A. stans [van] (3 sensu Choisy, Prodr. monogr. Hyperic.: 61 (1821).

Specimen in Herb. De Candolle (G-DC), not located.
A. cubense Griseb., Cat. PI. Cub.: 40 (1886); Sauvalle in An. Acad.
Cienc. med. fis. nat. Habana 5: 203 (1868), Fl. cub.: 8 (1869).
Type: Cuba, Pinar del Rio, 'juxta pineta pr. Pinales, 1860-1864',

127

Wright 2128 (GOET-holotype; BM!, GH!, K!, MO!, NY!, US!-
isotypes).

A. tetrapetalum (Lam.) Vail in Small, Fl. s.e. U.S.: 785 (1903), Man.
s.e.fl.: 868 (1933); Leon & Alain, Fl. Cuba 3: 315 (1953); W.P.
Adams in Rhodora 59: 93 (1957).

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 348, f.
164(1988).

Shrub or perennial herb woody at base, 0.2-1 m tall, erect, often
unbranched, branches divaricate or ascending. Stems green, (2)4-
lined and ancipitous when young, becoming 2-lined to terete; cortex
exfoliating in strips or flakes; bark thin, reddish brown, not corky.
Leaves sessile, (5-)8-35 x 4-15 mm, oblong-ovate or ovate to
triangular-ovate, margin subrecurved, paler beneath, not glaucous,
coriaceous, eventually deciduous at basal articulation, apex apiculate
or obtuse to rounded, base cordate-amplexicaul; venation: only one
pair of basal laterals visible; laminar glands dense, large; inframar-
ginal glands dense. Inflorescence l(3)-flowered with branching
pseudo-dichotomous, occasionally with short lateral branches from
up to 3 nodes below; pedicels 3-12 mm long; bracts foliar. Flowers
20-30 mm or more in diam.; buds compressed-subglobose. Sepals
4, markedly unequal, not? enlarging in fruit; outer7-13(-15) x 5.5-
9(-10) mm, broadly ovate, foliaceous, subapiculate to obtuse, base
cordate, basal veins 3(5), unbranched; inner 7-15 x 2-c. 3 mm,
narrowly lanceolate, acute, basal veins 1-3. Petals 4, bright yellow,
10-15 (or larger?) x 7-10 mm, 1.2-1.3 x sepals, obovate-oblong,
with apiculus lateral, acute to incurved, acuminate. Stamens c. 100,
longest c. 4.5-6.5, c. 0.45 x petals. Ovary 3-merous, 3-3.5 x 1.6-2
mm, pyramidal-ovoid to ellipsoid-ovoid, acute, placentation pari-
etal; styles 3, 3-3.5 mm, 1-1.2 x ovary, divergent distally. Capsule
c. 5-6 x 3.5^1, broadly ellipsoid-ovoid to subglobose, 3-lobed.
Seeds blackish brown, c. 0.7 mm long, ecarinate; testa shallowly
scalariform. 2n = 18 (n = 9, Adams in Robson & Adams, 1968).

Moist low pinelands and roadside ditches, on sandy soil; lowland.

U.S.A. (Florida and adjacent Georgia and Alabama), Cuba (Pinar
del Rio).

U.S.A. Alabama: Baldwin Co., E. side of US 90-98, E. from Mobile by
causeway, 7 June 1971 (fl), Krai 43061 (BM). Florida: Duval Co., 4.3 km S.
of Nassau Co., line on US route 1 and 23, NW of Jacksonville, 30 May 1973
(fr), Boufford 9265 (BM, MO*); Lake Co., Eustis, 16-30 June 1895, Nash
1977 (DAO*, K, MO*, NCU*, PH*); Sarasota Co., Fla 780, 25.1 km E. of
Sarasota, 20 June 1974 (fl), Baltzell 6501 (BM). Georgia: *Glynn Co.,
Brunswick, 5 April 1939 (fl & fr), Koelz 13466 (MO); *Irwin Co., 6.4 km N.
of Ocilla, Wilbur 3074 (FSU, NCSC, SMU); *McIntosh Co., Sapelo Island,
Duncan 17970(GA).

128

N.K.B. ROBSON

CUBA. Pinar del Rio: Vinales, La Vega, February 1867(fl), Wright 2\28
(BM, GH, MO, NY, S, US).

H. tetrapetalum is a derivative of 25. H. crux-andreae, differing from
it in the ovate to triangular-ovate leaves, which are strongly cordate-
amplexicaul and sometimes grade in form into that of the outer sepals,
although they are usually somewhat larger. The terminal pseudo-
dichotomous inflorescence is also nearly always diagnostic, but the
Alabama specimen cited is intermediate in this respect. These two
species overlap in distribution in southern Georgia and northern
Florida but do not otherwise intergrade morphologically.

27. Hypericum edisonianum (Small) W.P. Adams & N. Robson in
Rhodora 53: 1 5 ( 1 96 1 ); W.P. Adams in Contr. Gray Herb. Harv.
no. 189: 44 (1962), in J. Elisha Mitchell sclent. Soc. 89: 68
(1873); D.B. Ward, Rare & endangered biota of Florida 5.
Plants: 94 (1980); Godfrey & Woolen, Aquatic & wetland pis
s.e. U.S. Dicots: 340 (1981). Type: U.S.A., Florida, Highlands
Co., '21 miles [33.6 km] east of Arcadia', Hand 118 (NY-
holotype).

Fig. 18C, Map 16.

Ascyrum edisonianum Small, Man. s.e.fl.: 868 (1933); W.P. Adams
in Rhodora 59: 91 (1957).

Shrub 0.3-1 .5 m tall, erect, often unbranched below, much branched
pseudo-dichotomously above, forming spreading flat-topped crown;
horizontal roots bearing adventitious shoots form thickets. Stems
red-brown, 4-6-lined and ancipitous when young, soon 2-lined;
cortex exfoliating in strips; bark thin, reddish brown to grey, not
corky. Leaves sessile, appressed or ± spreading, 15-26 x 5-8(-

ll)mm, elliptic, margin subrecurved to subincrassate, paler be-
neath, markedly glaucous above only, coriaceous, soon deciduous at
base leaving prominent gland-like auricles, apex obtuse to acute,
base cuneate to subrounded; venation: up to 4 pairs of laterals
sometimes visible: laminar glands dense. Inflorescence 1 -flowered
with repeated pseudo-dichotomous branching; pedicels 3-5 mm;
bracts foliar; bracteoles lanceolate. Flowers 15-20 mm in diam.;
buds compressed-subglobose. Sepals 4, markedly unequal, not?
enlarging in fruit; outer (8-)9-13(-17) x (5-)6-8(-9) mm, broadly
ovate, acute to subacuminate, base cordate, basal veins 5-7, midrib
not or obscurely branched; inner c. 5-6 x 0.6-1.2 mm, linear-
lanceolate, acuminate, basal veins 5, midrib unbranched. Petals 4,
bright? yellow, (10-) 12-1 8 x (5-)6-l 1 mm, c. 1 .2 x sepals, obovate
with apiculus lateral, acute. Stamens c. 70-80, longest 6-7 mm, c.
0.5 x petals. Ovary 3^-merous, 3.5-4 x c. 1.5 mm, narrowly
triangular-ovoid, acute, placentation parietal?; styles 3-4, 2-3 mm
long, 0.6-0.75 x ovary, wholly appressed. Capsule 5-8 x (3?-)4
mm, triangular-ovoid, acute, 3-4-lobed. Seeds brown to yellow-
brown, c. 0.8 mm long, ecarinate; testa 'reticulate' (Godfrey &
Woolen).

In sandy soil of low prairies, in marshy areas in pine flatwoods and
al pond margins; lowland.

U.S.A. (cenlral peninsular Florida - Highlands, Glades and De Solo
Counlies).

U.S.A. Florida: Glades Co., Fla. 17, 9.6 km NW of Palmdale, 7 May
1 975 (fl & e. fr),Baltzell 7285 (BM, FLAS*); Highlands Co., 8 km NE of Old
Venus in Old State Road 8 (Childs to Venus), 30 September 1979 (fl), Judd
2517(BM, FLAS*).

H. edisonianum is a derivative of 25. H. crux-andreae, from which
il can be distinguished by the smaller, thicker, obtuse to acute leaves,
the pseudo-dicholomous branching and Ihe paired persislenl gland-
like auricles al Ihe base of each leaf.

Ward ( 1 980) suggested that H. edisonianum had been isolated on
the Lake Wales Ridge of cenlral peninsular Florida during Pleistocene
flooding, when much of Florida was benealh sea level.

28. Hypericum suffruticosum W.P. Adams & N. Robson in Rhodora
63: 15 (1961), sphalm. ' suffructicosurn '; W.P. Adams in Contr.
Gray Herb. Harv. no. 189: 49 (1962), in / Elisha Mitchell
sclent. Soc. 89: 68 ( 1 973); Radford, Ahles & Bell, Man. vase, pis

Map 16 Sect. 20: 27. H. edisonianum A specimens, A records; 28. H. suffruticosum
multicaule • specimens, O records, also in Massachusetts, Nantucket I.

specimens. D records; 29. H. hypericoides: b. subsp.

STUDIES IN HYPERICUM

129

Carolina*: 710 (1968); R.C. Clarke in Ann. Mo. hot. Gdn 58:
210 (1971); Clewell, Guide vase, pis Florida Panhandle: 372
(1985); Godfrey, Trees, shrubs & woody vines n. Florida, etc.:
353, f. 167 (1988). Type as for Ascyrum pumilum Michaux.
Fig. 18D, Map 16.

Ascyrum pumilum Michx., Fl. bor-amer. 2: 77 (1803); Pursh, Fl.
Amer. sept. 2: 373 (1814); Elliott, Sketch hot. S. Carolina 2: 21
(1821); Choisy, Prodr. monogr. Hyperic.: 60 (1821), in DC.,
Prodr. 1: 555 (1824); Spach, Hist. nat. veg. Phan. 5: 463 (1836),
in Annls Sci. nat. (Bot.) II, 5: 369 (1 836); Torrey & Gray, Fl. N.
Amer. 1: 1 56(1 838); Chapm., Fl. South. U.S.: 39 (1865); Coulter
in Bot. Gaz. 11: 79 (1886), in A. Gray, Syn. fl. N. Amer. 1: 283
(1897); Small, Man. s.e.fl.: 867 (1933); W.P. Adams \nRhodora
59: 77 ( 1 957); nonHypericumpumillum Sesse & Mocino ( 1 894).
Type: U.S.A., Georgia, without precise locality, Michaux s.n. (P-
holotype, BM!-microfiche).

A. pauciflorum Nutt., Gen. N. Amer. pis 2: 15(1818); Choisy, Prodr.
monogr. Hyperic.: 60 (1821), in DC., Prodr. 1: 555 (1824); non
Hypericum pauciflorum Kunth (1822). Type: U.S.A., Georgia, 'e
sylvestris Georgiae', Nuttall s.n. (PH-holotype; BM!).
A. nummularifolium Banks [in sched.] ex Steud., Nomencl. hot. 2nd
ed. : 77(1 840), nomen.

Icon: Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 353, f.
167(1988).

Q

Dwarf shrub 0.05-0. 15(-0.2) m tall, erect, few- to many-stemmed
and cushion-like or decumbent and ± matted, branched from near
base and sparingly pseudo-dichotomously, branches suberect or ±
spreading. Stems becoming red-brown, 4-lined and ancipitous when
young, soon narrowly 2-winged; cortex exfoliating in thin strips or
flakes; bark thin, reddish brown, not corky. Leaves sessile to
subsessile, ± spreading, (3-)4-8(-10) x (l-)2-3 mm, narrowly
obovate or oblanceolate to elliptic or oblong-linear, margin plane to
subrecurved, paler beneath, sometimes slightly glaucous,
subcoriaceous, eventually deciduous above base, apex obtuse, base
rounded to cuneate; venation: only midrib visible; laminar glands
dense; inframarginal glands dense. Inflorescence 1 -flowered, often
with a pair of pseudo-dichotomous branches; peduncle 0.5 mm long;
pedicel (5-)7-10(-12) mm long, mostly recurved to reflexed at
maturity; bracts foliar; bracteoles linear-subulate. Flowers c. 10-15
mm in diam.; buds compressed-subglobose. Sepals 4 and markedly
unequal or 2, enlarging somewhat in fruit; outer (4-)5-8 x (4-)4.5-
6(-8) mm, broadly ovate to broadly elliptic, obtuse to rounded, base
truncate to angustate, near-basal veins 3, all branched; inner (when
present) minute. Petals 4, pale yellow, (4-)5-8 x (4-)5-6(-8) mm,
often unequal, narrowly obovate, with apiculus terminal, obtuse,
fugaceous. Stamens c. 30, longest 2.5-4 mm, 0.4-0.6 x petals.
Ovary 2-merous, 2-2.5 x 0.6-1 .3 mm, narrowly compressed, cylin-
dric-ellipsoid, obtuse, placentation parietal; styles 2, 1.2-1.5 mm,
0.6 x ovary, appressed or divergent above. Capsule 3-5 x 2-3 mm,
cylindric-ellipsoid, rounded, compressed. Seeds c. 1 mm long,
scarcely carinate; testa finely reticulate. 2n = 18 (n = 9, Adams in
Robson & Adams, 1968).

Dry sandy open pinelands of the coastal plain: lowland.

U.S.A. (southern North Carolina to eastern Louisiana, excluding
most of peninsular Florida).

U.S.A. Alabama: *Covington Co., Hardin & Duncan 14974 (GA);
*Washington Co., Fruitdale, 5 May 1904 (fr), ? (MO). Florida: Bay Co., 3.2
km E. of Lynn Haven, 20 May 1961 (n & fr),Adams 774 (K); Columbia Co.,
Lake City, 1 1-19 July 1895 (fr), Nash 221 1 (GH*, K, MICH*. MO*, NY*,
US*); Duval Co., near Jacksonville, April (fl), Curtiss 246 (BM, F*, FLAS*,
GH*, K, MISSA*, MO*, NY*, PH*, US*). Georgia: Bulloch Co., S. of
Portal, c. 65 m, 6 May 1982 (fl & fr), Bouffordet al. 22814 (A*, BM, MO*);
Camden Co., S. of Kingsland, 12 May 1930 (fl & fr), Moldenke 1 180 (K,
MO*). Louisiana: *St. Tammany Par., Bougere 2006 (LSU). Mississippi:
*Harrison Co., Biloxi, Tracy 4489 (F, MICH, MO, NY, OS, US); *Jackson
Co., May 1930, Donald s.n. (MISSA). North Carolina: *Bladen Co., Wood
8499 (GH). South Carolina: *Colleton Co., 17.6 km NW of Walterboro,
Adams 84 (FSU); Jasper Co., US 601, c. 2.2 km N. of junction of Co. Route
27-169, 22 June 1968 (fl), Leonard & Radford 1678 (BM, H).

Adams ( 1 957) related H. suffruticosum to 29. H. hypericoides (using
Ascyrum nomenclature); but the link seems clearly to be with 25. H.
crux-andreae through the reduced form of that species that has been
named Ascyrum cuneifolium Chapman, which occurs in extreme
north-eastern Florida and Alabama. H. suffruticosum is thus derived
from a different part of//, crux-andreae from that which gave rise to
24. H. edisonianum (q.v.).

29. Hypericum hypericoides (L.) Cr., Inst. rei herb. 2: 520 (1776);
Adams & Robson in Rhodora 63: 15 (1961); W.P. Adams in
Contr. Gray Herb. Harv. no. 1 89: 46 (1962), in Rhodora 64: 237
(1962), in}. Elisha Mitchell scient. Soc. 89: 68 (1973); Radford,
Ahles & Bell, Man. vase, pis Carolinas: 710 (1968); R.C. Clark
in Ann. Mo. hot. Gdn 58: 209 ( 1 97 1 ); R. Long & Lakela, Fl. trop.
Florida: 606 (1971); N. Robson in Taxon 29: 272 (1980);
Godfrey & Wooten, Aquatic & wetland pis s.e. U.S. Dicots: 341
( 1 98 1 ); Clewell, Guide vase, pis Florida Panhandle: 372 ( 1 985);
Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 354, f.
168 (1988); N. Robson in Bull. nat. Hist. Mus. Lond. (Bot.) 23:
68 (1993), in Eur. Gdn Fl. 4: 70, ff. 10.7, 10.8, 10.13 (1995).
Type as for Ascyrum hypericoides.

Maps 16, 17.

Ascyrum hypericoides L., Sp. pi: 788 (1753) excl. syn. Hort. Cliff.
et Plukenet., 2nd ed.: 1 108 (1763) excl. syn. Plukenet.; Marshall,
Arbust. amer.: 14 (1785) ['hypericodes']; Willd., Sp. pi. 3: 1473
(1802); W.T. Aiton, Hort. kew. 2nd ed. 3: 430 (1812) pro pane;
Choisy, Prodr. monogr. Hyperic. : 6 1 ( 1 82 1 ), in DC., Prodr. 1: 555
(1824); Griseb. Fl. Br.W.I.:] 12 (1860); Coulter in Bot. Gaz. 11:
80 (1886), in A. Gray, Syn.fl. N. Amer. 1: 283 (1897); Small, Fl.
s.e. U.S.: 785 (1903), Man. s.e.fl.: 867 (1933); Urban, Symbol.
Antill. 4: 41 1 (1910); Engl. in Engl. & Prantl, Nat. Pflanzenfam.
2nd ed. 21: 174 (1925) pro parte, excl. syn. A. crux-andreae L.;
Fawcett & Rendle, Fl. Jamaica S: 202, f. 79 (1926); Leon &
Alain, Fl. Cuba 3: 315 (1953); W.P. Adams in Rhodora 59: 79
(1957), in op. cit.64: 237 (1962); Correll & Johnston, Man. vase,
pis Texas: 1060 (1970); R.G. & M.L. Brown, Woody pis Mary-
land: 233 (1972); Strausbaugh & Core, Fl. W. Virginia 2nd ed.:
638 (1980). Type: Hispaniola, Hypericoides frutescens erecta,
flare luteo Plumier, Nov. PI. Amer. Gen.: t. 7 (1703) (lectotype -
Robson, 1980: 272). Fawcett & Rendle (1926: 203) indicated the
type as Patrick Browne's specimen from Jamaica (LINN 944.2)
named Hypericum angustifolium in Solander's hand with correc-
tion toAsc\rum, the correct specific epithet having been added by
Sir J.E. Smith; and Adams ( 1 957: 83) designated this specimen as
a neotype, it having been added to the Linnaean herbarium only
in 1758, after the publication of Specie s plantarum (1753). As I
have explained, however (Robson, 1 980), there is no need for the

130

N.K.B. ROBSON

Map 17 Sect. 20: 29. H. hypericoides: a. subsp. hypericoides • specimens, O records, A extinct? (Illinois).

selection of a neotype, as Plumier's illustration is an appropriate

lectotype.
A. perforata Lam., Encycl. 1: 285 (1783), nom. illegit. (Art. 63.2)

[ 'Ascyre perforce'].
Hypericoides perforata Poir., Tab. encycl. 3: 201 , t. 644 f. 2 ( 1 823).

Shrub to wiry shrublet (0.05-)0. 1-1 .2(-l .5) m tall, erect or decum-
bent to prostrate, unbranched or with ± sparse lateral branches or
branched from base and ± diffuse, sometimes mat-forming, lateral
branches erect or ± spreading, sometimes also branched pseudo-
dichotomously. Stems red-brown, 2-winged, the subsidiary lines
only visible when very young; cortex exfoliating in thin strips or
flakes; bark thin, reddish brown, not corky. Leaves sessile, spreading
to rarely ascending, (5-)7-25(-34) x (1-)1 .5-6(-8.5) mm,
oblanceolate to narrowly oblong or linear, margin subrecurved,
slightly paler beneath, not glaucous, subcoriaceous, eventually de-
ciduous just above base, apex rounded to obtuse, base rounded or
usually cuneate with paired basal glandiform auricles; venation: 1-
2 pairs of basal or near-basal veins sometimes visible, tertiary
reticulation obscure or not visible; laminal glands dense. Inflores-
cence 1-3-flowered from 1-4 nodes, sometimes with flowering
branches from up to 10 nodes below, the whole narrowly cylindric to
narrowly pyramidal, or sometimes with a pair of inflorescences of
the above type developing from one pseudo-dichotomous branch-
ing, or branching wholly pseudo-dichotomous; peduncle + pedicel
3-6 mm long, pedicel up toe. 1 mm (i.e. bracteoles touching or very
near sepals); bracts foliar; bracteoles triangular-subulate. Flowers
10-15(-20) mm in diam.; buds compressed-globose to compressed-
ellipsoid. Sepals 4, markedly unequal, enlarging somewhat in fruit;
outer 5-12.5 x (2-)4-12(-13) mm, ovate-suborbicular to narrowly

elliptic, obtuse or subapiculate to acute, margin plane, base shallowly
cordate to broadly cuneate, basal veins 3, outer pair branched; inner
minute or obsolete, up to c. 4 x 2 mm, lanceolate, acute, basal veins
3, outer pair branched. Petals 4, bright to pale yellow, (6-)8- 1 1 (- 1 2)
x 2-4(-5) mm, 1(-1 .5) x outer sepals, obovate to narrowly oblong-
elliptic, apiculus subapical, obtuse (very short) or absent. Stamens c.
40-50, longest 3-5 mm long, c. 0.5 x petals. Ovary 2-merous, 3-4
x 0.8-1 .5 mm, narrowly ovoid, acute, placentation parietal; styles 2,
0.5-1 mm long, becoming outcurved. Capsule 5-9 x 2-4 mm,
narrowly compressed-ovoid to narrowly compressed-cylindric-el-
lipsoid, acute. Seeds purple-brown, 0.6-0.8 mm long, ecarinate;
testa finely linear-reticulate to linear-foveolate. 2n = 18 (n = 9,
Adams in Robson & Adams, 1968).

Dry open sandy habitats, especially pinewoods, roadsides and hill-
sides, or moist shady woods and thickets, bogs and lake margins;
lowland to 840 m (U.S.A., Arkansas), 1 800 m (Guatemala), 1 650 m
(Jamaica) or 2900 m (Dominican Republic).

U.S.A. (south-east of line: eastern Texas, eastern Oklahoma to
southern Missouri and eastward to New York (Long Island) and
Massachusetts), eastern Mexico, Guatemala, Honduras Republic,
Cuba, Hispaniola, Porto Rico, Jamaica, Bahamas, Bermuda. Intro-
duced into the Azores.

H. hypericoides is clearly related to 25. H. crux-andreae but can be
constantly distinguished by the 2-merous gynoecium, narrower
leaves, smaller flowers and more richly branched stems. It is very
variable in habit and in leaf and sepal shape and size. The form with
erect stems unbranched at the base, narrowly oblong leaves and
cylindric inflorescence (mainly in the S. Carolina/Georgia area) is

STUDIES IN HYPERICUM

morphologically nearest to H. crux-andreae. From this form, two
lines of evolution have both resulted in plants with diffuse to
prostrate stems, one in the northern range of the species in the U.S.A.
(subsp. multicaule) and the other at high altitudes in the Dominican
Republic (subsp. pmstratum). The former has been treated as a
species (H. stragulum Adams & Robson) and a variety (H.
hypericoides var. multicaule (Michx. ex Willd.) Fosberg), but sub-
species would seem to be the appropriate rank for this taxon.

29a. Hypericum hypericoides subsp. hypericoides

Map 17.

Hypericoides frutescens, erecta, flare luteo Plum., Nov. pi. amer.:

'51 (1703).
Ascyrumfoliis oblongis, fruticosum, minis erectis Burm., PL amer.:

146, t. 152f. 1 (1758).
Ascyrum crux-andreae [var.] p angustifolium Nutt., Gen. Amer. Pis

2: 1 6 ( 1 8 1 8); Choisy, Prodr. monogr. Hyperic. : 6 1 ( 1 82 1 ), in DC.,

Prodr. 1: 555 ( 1 824) pro parte, excl. syn. A. multicaule; Torrey &

Gray, Fl. N. Amer. 1: 156 (1838) ['angustifolia]. Type: U.S.A.,

'hab. in Carolina', Nuttall s.n. (PH?-holotype).
A. linifolium Spach, Hist. nat. veg. Phan. 5: 459 ( 1 836), in Annls Sci.

nat. (Bot.) II, 5: 368 (1836); Britton & Millsp., Bahama fl.: 280

( 1920).Type: U.S.A., Louisiana, Covington, 1 832 (fl), Drummond

s.n. (P-holotype; BM!).
A. michauxii Spach, Hist. nat. veg. Phan. 5: 460 ( 1 836), in Annls Sci.

nat. (Bot.) II, 5: 368 (1836). Type: U.S.A., 'Ascyrum multicaule

var.' Michaux (P-holotype). In his second paper, Spach cites his

synonym as 'A., amplexicaule var. Michx.! herb.'. I have not seen

a specimen.
A. oblongifolium Spach, Hist. nat. veg. Phan. 5: 461 (1836), mAnnls

Sci. nat. (Bot.) II, 5: 369 (1836). Type as for A. crux-andreae var.

angustifolium Nutt.

A. montanum Raf., Fl. Tellur.: 1 16 (1838). Type not located.
A. plumieri Bertol. in Memorie Accad. Sci. Bologna 4: 77 (1853),

Misc. hot. 12: 19, t. 3 f. 3f, g ['plumierii^.Type: U.S.A., Alabama,

Gates s.n. (BO-holotype, BM!-microfiche).
A. crux-andreae sensu Chapm., Fl. South. U.S.: 38 (1865); Griseb.,

Cat. PI. Cub.: 40 (1866); Sauvalle, Fl. cub.: 8 (1868).
A. macrosepalum S. Brown in Jl N.Y. hot. Gdn 13: 192 (1912);

Britton, Fl. Bermuda: 245 & t. (1918). Type; Bermuda, Paget

Marsh, 29 November- 14 December 1912 (fr), Brown & Britton

1 136 (NY!-holotype; F!, GH!, MO!, PH, US!-isotypes).
A. hypericoides var. typicum Fernald in Rhodora 38: 432 (1936).

Type as for A. hypericoides L.
A. hypericoides var. oblongifolium (Spach) Fernald in Rhodora 38:

433 (1936), in Gray's Man. Bot.: 1008 (1950); R. A. Vines, Trees,

shrubs & woody vines ofS.W.: 754 (1960).
Hypericum hypericoides (L.) Cr. var. hypericoides, R. Long &

Lakela, Fl. trop. Florida: 606 (1971).

Icones: Mohlenbr., ///. Fl. Illinois fl. pis Hollies to Loasas: 18, f. 5
(1978); Godfrey, Trees, shrubs & woody vines n. Florida, etc.: 354,
f. 168(1988).

Plant erect, with main stem usually unbranched from base, 0.3-1 .5
m tall, freely branched well above ground level. Leaves oblong to
linear (broadest at the middle), 7-25 x 1-8.5 mm. Inflorescence-
branching dichasial/lateral to pseudo-dichotomous.

Habitats of the species, but absent in the the Dominican Republic
from above 2000 m.

Distribution of the species except for absence north of the line:
Virginia, Kentucky, southern Missouri, Oklahoma.

131

U.S.A. Alabama: Dallas Co., 8 km E. of Selma on Dallas road, c. 43 m,
23 September 1965 (fr), Demaree 52975 (BM); Lee Co., Auburn, October
1 896 (fr), Earle & Baker s.n. (Z). Arkansas: Montgomery Co., Mt Ida, S. fork
ofOuachitaR.,210m, 1 October 1 962 (fr), Demaree 46620 (BM); Pope Co.,
Nogo, 11 September 1932 (fr), Merrill 27 (NY). Florida: Hendry Co.,
Labelle, 19 April 1930 (fl), Moldenke 11019 (BM, K, MO*); Wakulla Co.,
1 .6 km N. of St Mark's, 17 October 1958 (fr), Godfrey 57865 (BM, FSU*).
Georgia: Clarke Co., Barnet Shoales, 12 April 1925 (st), Maguire s.n. (BM,
GA*); Ogelthorpe Co., Lexington, 201 m, 25 September 1965 (fr), Demaree
53008 (BM). Illinois: *Hancock Co., Augusta, July 1842, Mead s.n. (MO)
(fide Mohlenbrock & Evans, 1972: 145), extinct? Kentucky: *Hickman Co.,
Bayou du Chien, 23 August 1923 (fr), McFarland 2 1 3 (GH, MO); *Marion
Co., 1 .6-2.4 km SE of New Market, 8 July 1939 (fl), Wharton 4686 (MO).
Louisiana: *Caddo Par., Shinners 21177 (GA, SMU): St. Tammany Par.,
Covington, 1832 (fl), Drummond s.n. (BM, K). Maryland: Dorchester Co.,
Baltimore, Blackwater National Wildlife Refuge, 20 August 1972 (e. fr),
Windier, Kuser & Shastny 4 1 30 (H); *Worcester Co., Fernald, Long & Fogg
5573 (GH, PENN). Mississippi: Harrison Co., Biloxi, 8 July 1900 (fl), Tracy
6897 (BM); *Jackson Co., Ocean Springs, 14 September 1891 (fr), Seymour
91914 (DUKE, F, GH, MO, NCU, SMU, TEX). Missouri: *Butler Co., 19
October 1905 (fr), Bush 3762 (GH, MO, US); *Dunklin Co., Campbell, 2
September 1910 (fr), Bush 6236 (MO). New Jersey: *Cape May Co., S.
Seaville to S. Dennisville, 8 August 1937 (fr), Langheim s.n. (MO). North
Carolina: Chowan Co., 9.6 km E. of St John's Indian Trail Road, 14 October
1 958 (e. fr), Ahles 5 10 1 1 (H); Orange Co., W. of Chapel Hill, c. 2.25 km from
NC54 on Orange Grove road, 3 1 August 1 973 (fr), Boufford 1 1 764 (A*, BM).
Oklahoma: *Atoka Co., Hopkins 2850 (OKL); Le Flore Co., Page, W. end of
Rich Mtn, Ouachita National Forest, 26 October 1966 (fr), Demaree 53254
(BM); * Wagoner Co., near Wagoner, 3 September 1913 (fr), Stevens 2602
(MO). South Carolina: *Berkeley Co., 16 km NE of Monck's Corners, 13
September 1 939 (fr), Godfrey & Tryon 82 1 7 (DUKE, F, GH, MO, NY, PENN,
TENN, US); Chesterfield Co., 9.6 km E. of Patrick, 1 1 August 1956 (fr),
Radford 1 5807 (K).Tennessee: *Coffee Co., Norris & Sharp 16269 (TENN);
*FranklinCo.,Hungland,6August 1 939 (fr), Svenson 10404 (MO); *Shelby
Co., near Memphis, 18 May 1920 (si), Palmer 17532(MO).Texas: * Anderson
Co., Wilcox near Balestinge, 1 August 1943 (fr), Barkley 13598 (MO);
Brazoria Co., Columbia, 24 March 1 9 14 (fl), Palmer 5006 (K); Wood Co., 6.4
km E. of Mineola, 10 August 1967 (fl), Mears & Kukkonen 2407 (H).
Virginia: *MathewsCo., 'Fort Nonsense', S. of Soles, 1 September 1925 (fl),
Wherry & Pennell 1261 1 (MO); Princess Anne Co., Creed's, 9 September
1935 (fr), Fernald, Long & Fogg 4941 (K).

MEXICO. Chiapas: 54.6 km SE of Comitan, Los Lagos, 1500 m, 18
January 1952 (fl), Carlson 2250 (MEXU, MICH); mun. Tenejapa, Colonia
San Antonio, 21 15 m, 8 August 1964 (fr), Breedlove 7050 (BM, F, MICH).
Hidalgo: Km. 284 on highway NE of Jacala, Puerto de la Zorre, c. 1500 m, 8
July 1948 (fl), Moore & Wood 3785 (A, MICH): Distr. Zacualtipan, mun.

132

N.K.B. ROBSON

Zacualtipan, mountains above Tzincuatlan, 1900 m, 7 November 1946 (fl &
fr), Moore 1880 (GH). Michoacan: Morelia, Laguna, 2000 m, Arsene s.n.
(JE). Nuevo Leon: Mun. Montemorelos, La Trinidad, 19 August 1939 (fl),
Muller 2851 (GH, MICH). Oaxaca: Cordillera, 1200-1650 m, 1840 (fr),
Galeotti 920 (K, W). Puebla: Huauchinango, near Catalina, 1650 m, 2
October 1944(fl & fr),Sharp44\ 139(GH);Zacapoaxtla, 13 September 1941
(fl & fr), Miranda 3290 (MEXU). San Luis Potosi: 20 km SE of Zargoza,
Mesita de Gallos, 2000 m, 8 July 1 955 (fr), Rzedowski 6090 (MEXU, MICH,
US).Tamaulipas: mun. Aldama, c. 40 km NNW ofAldama, 23 July 1957 (fl),
Dressier 1976 (GH, MEXU, MICH, MO); vicin. Marmolejo, 12August 1930
(fl). Bartlett 10908 (MICH). Vera Cruz: near Jalapa, 1200 m, 21 June 1901
(fl), Pringle 8515 (BM. C. F, GH, K. MEXU, MO, P, US, W, Z); mun.
Chiconquiaco.Guacamaya, 1900m, 1 1 May 1973 (fl), Ventura 8275 (MICH).

GUATEMALA. Alta Verapaz: Coban, 1290 m. May 1886 (fr), von
Turckheim 88 (GH, JE, K, US, Z); near San Cristobal Verapaz, 1300 m, 6
January 1973 (fl), Williams. Molina & Williams 42219a (F). Baja Verapaz:
Sierra de las Minas, near La Union, 1800 m, 18 January 1974 (fl), Williams
et al. 43556 (BM). El Quiche: between Chalul and Cotzal, 1800 m, 6
February 1946 (fl). Sharp 4679 (F). Huehuetenango: Trinidad, 13 August
1896 (fl), Seler 3083 (GH, US). Jalapa: between Guisiltepeque and Potrero
Carillo, 1800 m, 11 December 1939 (fr), Steyermark 33086 (F). Zacapa:
Sierra de las Minas, between Rio Hondo and Finca Alejandria, 1 000- 1 500 m,
1 1 October 1939 (fl), Steyermark 20726 (F).

HONDURAS REPUBLIC. Copan: between San Isidro and San
Cristobal, c. 16 km S. of Copan Ruinas, 26 August 1975 (fr), A. & A. Molina
30697 (MO). Intibuca: vicin. La Esperanza and Intibuca, 1500-1600 m, 31
January- 12 February 1950 (fl), Standley 25344 (F, US); Kms 9-1 1 between
La Esperanza and Marcala, 1600 m, 5 April 1956 (fl), A. & A. Molina 24290
(F, MO). La Paz: 20 km from Marcala, La Chorrera, 1 200 m, 24 March 1969
(fl), A. & A. Molina 24410 (F).

BERMUDA. Pembroke Marshes, 1 1 August 1913 (fl), Collins 232 (F,
GH, K, US, W); North Shore, 31 August-20 September 1905 (e. fr), Brown
& Britton 10 (A, F, K, US).

BAHAMAS. Andros: 1.6 km W. of main road, just N. of Love Hill, 7
June 1975 (fl), Hill 3136 (NY). Grand Bahama: Eight Mile Rocks, 5-13
February 1905 (o. fr), Britton & Millspaugh 2380 (F, K, US). Great Abaco: by
end of Norman Castle road, near Winding Bay Pond, 7 July 1 969 (fr), Proctor
30665 (BM). New Providence: S. of Delaporte Caves on Blake Road, 9 July
1960 (fl), Webster, Samuel & Williams 10373 (F, GH, US).

CUBA. Pinar del Rio: Arroya del Sumidero, 7 & 9 August 1912 (fl),
Shafer & Leon 13616 (BM, F, MO, US); Sabanalamar, El Sabalo, April 1950
(fl), Bro. Alain 1323 (G, H, US). Las Villas: Trinidad Mts, San Bias to Buenos
Aires, Gavinas, 18 September 1941 (fl), Gonzales 154(BM); nearManacas,
1 1 July 1 936 (f\),Smith & Hodgdon 3096 (US). Oriente: Sierra Maestra supra
Daiguiri, c. 1000 m, 29 October 1916 (fl), Ekman III 8172 (F, MICH, US):
Santiago, Gran Piedra, April 1949 (fl), Bro. Clemente 6496 (GH, US).

HAITI. Massif de la Selle, Morne des Commissaires, near Marc Etabli,
1500 m, 7 December 1941 (fl & fr), Holdridge 878 (BM, MICH, MO, NY,
US); vicin. of Mission, Fondo Varettes, c. 1000m, 17 April -4 May 1920(fl
& fr), Leonard 3828 (GH, NY, US).

DOMINICAN REPUBLIC. Barahona: between Pedernales andAceitial,
8-12 August 1946 (fl), R. & E. Howard 8160 (BM, GH, NY, US); Sierra de
Bearuco, near Canote, 15 January 1970 (fl), Terborgh 134 (A). La Vega:
prope Jarabacoa, 600 m, June 1912 (fr), Fuertes 1618 (GH, NY, US, W, Z);
5 kmW. of LaCulata (de Constanza), 1470-1500 m, February 1982 (fl & fr),
Zanoni et al. 19250 (NY). Monte Cristi: Sabaneta, Lagunas de Cenobf, 15
August 1 929 (fl), Vale ur 1 6 (US). Santiago Rodrfguez: San Jose de las Matas,
180-210 m, 22 April 1931 (fl), Valeur 754 (F, MICH, MO, NY, US);
Cabirmar, arroya Los Guanos, 8-4 km al SE de Los Ramones en el camino
hacia Manaclas, 630 m, 16 July 1985 Mejia, Pimentel & Garcia 1466 (BM,
JBSD*).

PUERTO RICO. Prope Utuado in montibus ad Cazuco, 9 March 1887
(fl), Sintenis 6377 (BM, F, GH, K, MO, US, W, Z); Laguna Tortuguero, 17
January 1968 (fl & fr), Howard & Nevling 16952 (A).

JAMAICA. Portland: c. 100-200 m W. of Silver Hill Gap, 1050 m, 10
July 1967 (fl & fr), R. & S. Weaver 948 (GH). St Andrews: near Bellevue, c.
1200m, 17 November 1957 (fl), Yuncker 17427 (BM, F, MICH). St Thomas:
Blue Mts, between Portland Gap and Blue Mountain Peak, 1 650-2240 m, 1 8
August 1954 (fl), Webster & Wilson 5458 (A, BM, MICH).

AZORES (introduced?). Faial: Coastal Branco, 8 October 1962 (fl),
Goncalves 740 (BM).

Subsp. hypericoides varies in leaf-width, the narrow-leaved form
(Ascwum linifolium) being found in the southern States from Loui-
siana to Florida as well as in the Bahamas and Bermuda (A.
macrosepalum). Britton (1918) suggested that it had been carried to
Bermuda by wind or birds from Florida and had there speciated, but
the Bermudan plants do not differ from other narrow-leaved forms.
The occurrence of this subspecies in the Azores could also be the
result of long-distance transport, but it could equally have been
introduced by man.

29b. Hypericum hypericoides subsp. multicaule (Michx. ex Willd.)
N. Robson in Taxon 29: 273 ( 1 980); in Cullen et al., Eur. Gdn Fl
4: 70 (1995). Type as for Ascyrum multicaule Michx. ex Willd.

Map 16.

Ascyrum multicaule Michx. ex Willd., Sp. pi. 3: 1472 (1802);
Michx., Fl. bor.-amer. 2: 77 (1803). Type: 'Hab. in Virginia,
Carolina', Michaux s.n. (extreme r.h. specimen) (P-holotype,
BM!-microfiche, GH-photograph).

A. crux-andreae [var.] P sensu Choisy, Prodr. monogr. Hyperic.: 61
(1821) pro parte, quoad syn. Ascyrum multicaule.

A. helianthemifolium Spach, Hist. nat. veg. Phan. 5: 460 (1836), in
Annls Sci. nat. (Bot.) II, 5: 368 (1836). Type: U.S.A., Louisiana,
Michauxl (P-holotype).

A. spathulatum Spach, Hist. nat. veg. Phan. 5: 462 (1836), in Annls
Sci. nat. (Bot.) II, 5: 369 (1836). Type: U.S.A., without precise
locality or collector (P-holotype, BM!- microfiche).

A. crux-andreae sensu Coulter in Bot. Gaz. 11: 80 (1886).

A. hypericoides var. multicaule (Michx. ex Willd.) Fernald in Rhodora
38: 433 ( 1 936), in Gray 's Man. Bot. : 1 008 ( 1 950); W.P. Adams in
Rhodora 59: 87 (1957); R.A. Vines, Trees, shrubs & woody vines
ofS.W.: 754 (1960); Correll & Johnston, Man. vase, pis Texas:
1080 (1970); Cooperrider in Castanea 54: 7, f. 1 (1989).

Hypericum stragulum Adams & Robson in Rhodora 63: 15 (1961);
W.P. Adams in Rhodora 64: 236 (1962), in J. Elisha Mitchell
scient. Soc. 89: 68 (1973); Radford, Ahles & Bell, Man. vase, pis
Carolinas: 711 (1968) ['stragalum']; nee H. multicaule Lam.
(1797) nee H. spathulatum (Spach) Steud. (1840). Type as for
Ascyrum multicaule Michx. ex Willd.

H. hypericoides var. multicaule (Michx. ex Willd.) Fosberg in
Castanea 30: 202(1965).

H. hypericoides var. multicaule (Michx. ex Willd.) Waterfall in
Rhodora 73:553(1911).

Icon: Mohlenbr., ///. Fl. Illinois fl. pis Hollies to Loasas: 19, f. 6
(1978).

Plant decumbent, with several to many ± branched stems, 0.05-0.3

STUDIES IN HYPERICUM

m long, forming mats. Leaves usually oblanceolate (broadest above
middle), 10-20 x 3-6 mm. Inflorescence-branching dichasial (1-3-
flowered) or lateral.

Dry slopes, road embankments and dry to moist rich woods; 70-
840m.

Eastern United States (Massachusetts - Nantucket I. to N. Carolina)
westward to south-eastern Kansas, Oklahoma and Texas.

U.S.A. Alabama: Henry Co., Mt Dale, n.d. (fr),Lecis s.n. (BM); Jackson
Co., Bryant, 3 July 1938 (fl), Porter s.n. (BM). Arkansas: Montgomery Co.,
Ouachita National Forest, Mt Ida, 210 m, 5 October 1960 (fr), Demaree
42916 (BM); Pope Co., Nogo, 1 1 September 1932 (fr), Merrill 27 (NY).
Delaware: New Castle Co., Townsend, 12 August 191 1 (fl), Churchill s.n.
(MO); *Sussex Co., Long & Bartram 1572 (PH). Distr. of Columbia:
Washington, July 1897 (fl). Holm s.n. (H); *Eckington, 14 July 1913 (fr),
Painter s.n. (MO). Georgia: Fulton Co., College Park, 300 m, 8 August 1961
(fl), Schallert 850 (BM); Whitfield Co., Dalton, 255 m, 10 August 1900 (fr),
Harper 394 (BM, K). Illinois: ? Co., southern Illinois, 1866 (fr), Vasey s.n.
(BM); *Pope Co., Golconda, 7 October 1919 (fr), Palmer 16698 (GH, MO,
NY, PH). Indiana: *Clark Co., Deam 5414 (IND, NY). Kansas: *Cherokee
Co., 7 May 1 897 (st), Hitchcock 1012 (GH, MICH, MO, NY, US). Kentucky:
*Bell Co., mountains around Pineville, August-September 1914 (fr), Mac-
kenzie 922 (MO, NY). Louisiana: *Caddo Par., July 1909, Cocks s.n. (NO).
Maryland: *Ann Arundel Co., Bartlett 1838 (MICH); *Calvert Co., S. of
Parker's Creek, 4 August 1956 (fl), Seymour 16854 (MO). Massachusetts:
*Nantucket Co., Nantucket I., Pennell 1 1 1 74 (PH). Mississippi: *PanolaCo.,
18 April 1898 (fl), Eggert s.n. (MO); *Union Co., 23 September 1891,
Seymour s.n. (DUKE). Missouri: Jasper Co., near Prosperity, Center Creek,
10 October 1923 (fl & fr), Palmer 24099 (MO*, Z); Jefferson Co., near Big
River, NNE of confluence with Parker Creek, 10.5 km W. of De Soto, c. 600
m, 20 July 1985 (fl), P.H. & T.E. Raven 26758 (BM, MO*). New Jersey:
Atlantic Co., Mays Landing, 31 August 1937 (fr), Moldenke 10155 (BM);
Burlington Co., c. 2.9 km S. of Chatsworth, 20 August 1948 (fl & fr),
Lawrence & Dress in Bailey 558 (BM). New York: *Nassau Co., Ferguson
7798 (GH). North Carolina: Macon Co., Highlands, Ravenel Lake, 27 August
1966 (fl), Bozemann 7970 (BM, H); Vance Co., Ruin Creek on U.S. 1 584, c.
6.4km SWof Henderson, 21 July l956(fr),Ahles 17322 (H). Ohio: * Adams
Co., 23 September 1923, Roads s.n. (OS); *Hocking Co., Laurel Twp., SW of
Logan, 2 August 1982 (fl), Lammers 4815 (MO). Oklahoma: *Carter Co.,
Sandy Canyon, 9.6 km E. of Ardmore, 31 October 1942 (fr), Hopkins 6340
(MO); *Garvin Co., Washita R., near Davis, 29 July 1933 (fl). Palmer 42045
(MO). Pennsylvania: *Chester Co., French Creek, July 190- (fr), Eby s.n.
(MO); *Lancaster Co., Susquehanna R., 22 August \S6\, Porter s.n. (CM);
*York Co., A dams 4380 (GH). Tennessee: *Knox Co., Knoxville, July 1 895
(fr), Ruth s.n. (F, GH, MO); Putnam Co., by T-40c. 8 kmW. of Cookeville and
junction Tenn. 42, 7 October 1970 (fl & fr), Krai 41437 (BM). Texas:
*Henderson Co., Eustace, Lundell & Lundell 9574 (GH, MICH, NY, SMU);
*Tarrant Co., above Bear Creek, W. of Irving, 7 July 1945 (fl), Lundell 14030
(MO). Virginia: *Wise Co., Big Stone Gap, 24 July 1891 (fl), Seymour 91724
(MO); Norfolk Co., prope Portsmouth, July 1840 (fl), Rugel 199 (BM). West
Virginia: *Cabell Co., Gilbert 548 (DUKE, F, PENN, PH, SMU, TENN,
WVA).

29c. Hypericum hypericoides subsp. prostratum N. Robson in
Bull. not. Hist. Mus. Land. (Bot.) 23: 68 (1993). Type: Domini-
can Republic, San Juan, Sabana Nueva, Cordillera Central N. of
Rio Arriba del Norte, 1950 m, 17-20 September 1944, R.A. &
E.S. Howard 9080 (BM!-holotype; GH!, MICH!, NY!, US!-
isotypes).

Hypericoides frutescens, humi-fusa, flare luteo Plum., Nov. pi.

amen: 52(1703).

Ascyrum foliis lanceolato-linearibus, higlandulis, ramis diffusis
Burm., PL amer.: 146, t. 152 f. 2 (1758).

Icon: Burm., PL amer.: t. 152 f. 2 (1758).

Plant prostrate, with stems ± numerous, radiating and branching,

133

forming mats. Leaves narrowly oblong to oblong-spathulate, 3-8(-
10) x 1-2.5 mm. Inflorescence-branching pseudo-dichotomous.

Open Pinus forest, grassland and open slopes; (1600-)! 800-2900

Dominican Republic (La Vega, Santiago, San Juan, Peravia).

DOMINICAN REPUBLIC. La Vega: Constanza, Ci^naga de la Culata,
1 500- 1 600 m, 28 November 1 969 (fr), Liogier 1 7052 (GH, NY, US); vicinity
of La Lagunita, 2800-2900 m, 1 9 July 1 967 (fl ), Gastony, Jones & Norris 3 1 7
(GH, NY, US). San Juan: Baoruco, Alto de Toro, 2100 m, 26-27 June 1973
(fl), A. & L Liogier 19751 (NY). Santiago: San Jose de Ocoa, La Horma
Arriba, 1900-2000 m, 1 May 1972 (fr), Liogier 18589 (F, NY, US); La
Rusilla, 2072 m, 26 March 1964 (fl), Jimenez 4918 (US). Peravia: 42 km al
NW de San Jose de Ocoa, nacimiento del Rio Las Cuevas en la bas del Monte
Tetero de Mejfa, 1940 m, 30 May 1984 (fr), Mejfa, Pimentel & Garcia 610
(BM, JBSD*).

Subsp. prostratum is clearly a higher-altitude derivative of the form
of subsp. hypericoides that occurs at lower levels in the Dominican
Republic. Although there is a region between 1600 and 2000 m
where both subspecies are present, there appears to be little morpho-
logical overlap between them.

Some specimens of subsp. prostratum have been determined as//.
constanzae Urban, but this is a synonym of H. diosmoides Griseb.
(sect. 29. Brathys), see Robson (1990: 37).

Sect. 21. WEBBIA (Spach) R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 3(6): 21 1 (1893).

Shrubs or sometimes single-stemmed and arborescent, up to 4 m
tall, deciduous, glabrous, without dark glands; branching lateral.
Stems 4-lined and ± compressed (ancipitous) when young, not
usually becoming terete in first season, ± glandular, especially on
lines; cortex dull reddish brown; bark smooth, pale reddish brown
to whitish grey. Leaves opposite or abnormally 3-whorled, decus-
sate, sessile, free, deciduous at basal articulation; lamina entire,
with venation pinnate, closed, with tertiary reticulation dense;
laminar glands punctiform; marginal gland dots dense, relatively
large; ventral glands absent. Inflorescence up to 30-flowered, with
branching dichasial/monochasial from 1-5 nodes, sometimes with
flowering branches from up to 7 lower nodes; bracts and bracteoles
foliar or ± reduced. Flowers stellate, homostylous. Sepals 5, partly
united, persistent, erect to spreading or subreflexed in fruit, with
margin glandular; veins 5-7; laminar glands linear to punctiform;
marginal glands very small, sessile or on short cilia; submarginal
and inframarginal glands absent. Petals 5, persistent, spreading
and twisting after flowering, with apiculus obsolete or absent;
margin entire; marginal glands absent; laminar glands linear. Sta-
men fascicles 3 (i.e. united 2+2+1), distinct, with stamens 36-75;
filaments basally united; anthers yellow, gland amber; pollen type
L Ovary with 3 (sometimes incompletely) axile placentae, °o-
ovulate; styles 3, free, bases divaricate; stigmas subclavate to
narrowly capitate. Capsule 3-valves, coriaceous, with valves finely
and obscurely striate. Seeds narrowly cylindric or conico-cylindric,
not or slightly carinate, apically truncate; testa linear-reticulate to
linear-foveolate.

BASIC CHROMOSOME NUMBER (x). 10;ploidy4.

134

N.K.B. ROBSON

HABITAT. Evergreen Laurus forest, secondary growth after defor-
estation, and dry bushland on rocky ground; (20-) 180- 1200 m.

DISTRIBUTION. Canary Islands (all islands except Lanzarote and
(probably) Fuerteventura), Madeira.

1 species.

1. Hypericum canariense L., Sp. pi.: 784 (1753); Miller, Card.
Diet. 8th ed. no. 4 corrig. (1768); Lam., Encycl. 4: 155 (1797);
Willd., Sp. pi. 3: 1448 ( 1 802); Buch in Abh. K.Akad. Wiss. Berlin,
Phys. Kl. 1816-1817: 366, 371, 380 (1817); Choisy, Prodr.
monogr. Hyperic.: 40 (1821), in DC., Prodr. 1: 544 (1824);
Trevir., Hyper, sp. animadv.: 8 (1861); Masferrer in An. Soc. esp.
Hist, nat 9: 28 (1880); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 3(6): 211(1 893), 2nd ed. 21 : 1 76 ( 1 925); Stefanoff
in God. Agr.-les. Fak. Univ. Sofiya 10: t. 2 f. 7 (1932), 11: 145
(1933), 12: 82 (1934), in Pflanzenareale 4: Karte 2a (1933); Lid
in Skr. norske Vidensk.-Akad. I Math.-Nat. Kl., N.S. No. 23: 1 19
(1967); Schaeffer, PL Canary Is.: 146, 147 (photograph) (1967);
Kunkel in Mitt. dt. dendrol. Ges. 65: 108 (1972); Voggenr. in
Dissnes. hot. 26: 655, 688 ( 1 974); Kunkel, Endemismos Canarios:
29 1 ( 1 977); Bramwell, D. & Z.J., Wildfls Canary Is. : 1 62, f. 1 98
( 1 984); I. Hagemann in Flora 183: 242-252, ff. 1 3-20 ( 1 989); N.
Robson in Cullen et al., Eur. Gdn Fl. 4: 70, ff. 1 1 . 1 , 1 1 . 1 1 ( 1 995).
Type: Cult, ex Canary Is., 'Crescit in insulis Canariis', Hon.
Cliff.: 38 1 , no. 9 (BM!- lectotype; seeWijnands, Bot. Commelins:
109, 1983).

Fig. 19, Map 18.

H. floribun dum Aiton, Hort. Kew. 3: 104 (1789); Lam., Encycl. 4:

155 (1797); Willd., Sp. pi. 3: 1448 (1802); Buch in Abh. K.Akad.
Wiss. Berlin, Phys. Kl. 1816-1817: 372, 380 (1817); Choisy,
Prodr. monogr. Hyperic. : 40 ( 1 82 1 ), in DC., Prodr. 1 : 544 ( 1 824);
Rchb., Ic. hot. exot. 1: 64, t. 95 (1827); Link in Buch, Phys.
Beschr. Canar. Ins.: 153, 167 (1828); Lowe, Fl. Madeira: 76
(1868); Masferrer in An. Soc. esp. Hist. nat. 9: 28 (1880);
Menezes, Fl. Archip. Madeira: 29 (1914); R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 3(6): 211 (1893), 2nd ed. 21: 177
(1925); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: tt. 1 f. 3,
3 f. 9 (1932), 11: 145 (1933), 12: 82 (1934); Grabham, Pis seen
in Madeira: 94 (1934); Lid in Skr. norske Vidensk-Akad., Math.-
Nat. Kl., no. 23: 120 (1967); Kunkel, Endemismos Canarios: 293
(1977). Type: Madeira, 1777 (fl), Masson s.n. (BM!-holotype &
isotype).

H. debile Salisb., Prodr. stirp. horto Chapel Allerton: 396 (1796),
nom. illegit. (Art. 63.1). Type as for H. canariense L.

H. corymbosum Moench, Methodus, Suppl.: 41(1 802), nom. illegit.
(Art. 63.1). Type as for H. canariense L.

H. canariense var. montanum Buch in Abh. K. Akad. Wiss. Berlin,
Phys. Kl. 1816-1817: 382 (1817) ['montana'], nomen.

H. canariense var. [p] triphyllum Choisy in DC., Prodr. 1: 544
(1824). Type not found.

H. canariense var. [y] salicifolium Choisy in DC., Prodr. 1: 544
(1824). Type not found.

Webbia floribunda (Aiton) Spach, Hist. nat. veg, Phan. 5: 409
( 1 836), in Annls Sci. nat. (Bot.) II, 5: 356 ( 1 836); Webb & Berth.,
Phytogr. canar. 1: 47, t. 4B (1836).

W. heterophylla Spach, Hist, nat veg. Phan. 5: 409 (1836). Type:
'Hypericum canariense Linn.?'.

28-

17

16

Map 18 Sect. 21: 1 . H. canariense • specimens, O record. Limits on Tenerife according to Voggenreiter ( 1974).

STUDIES IN HYPERICUM

W. platypetala Spach, Hist. nat. veg. Phan. 5:410(1 836). Type: cult.

in horto Paris. (P-holotype). This is an error for platysepala, see

below.
W. canariensis (L.) Webb & Berth., Phytogr. canar. 1: 48, t. 4C

(1836); Pitard & Proust, Les lies Canaries: 133 (1909).
W. platysepala Spach in Annls Sci. nat. (Bot.) II, 5: 356 (1836);

Webb & Berth., Phytogr. canar. 1: 49, t. 4D ( 1 836). Type as for W.

platypetala Spach.
Hype ricum platypetalum (Spach) Steud., Nomencl. bat. 2nd ed. 1:

789(1840).
H. platysepalum (Spach) Walp., Repert. hot. syst. 1: 386 (1842);

Masferrer in An. Soc. esp. Hist. nat. 9:28(1 880).
H. canariense [van] a typicum Bornm. in Bot. Jb. 33: 452 (1903)

['rypic-fl']; Ceballos & Ortufio, Veg. Fl. for. Canar. occid.: 388

(1951). Type as for H. canariense L.
H. canariense [var.] fifloribundum (Aiton) Bornm. in Bot. Jb. 33:

453 (1903) ['floribunda']; Ceballos & Ortuno, Veg. Fl.for. Canar.

occid.: 388(1951).
H. canariense [var.] y platysepalum (Spach) Bornm. in Bot. Jb. 33:

453 (1903) ['platysepala}.
Wehbia canariensis var. [a] typica (Bornm.) Pitard & Proust, Les

lies Canaries: 134(1909).
W. canariensis [var.] ft floribunda (Aiton) Pitard & Proust, Les lies

Canaries: 134(1909).
W. canariensis [var.] y platypetala (Spach) Pitard & Proust, Les lies

Canaries: 134(1909).

Icones: Cooke in Bot. Cabinet 10: t. 953 ( 1 824); Rchb., Ic. hot. exot.
1: t. 95 (1827); Webb & Berth., Phytogr. canar. 1: tt. 4B-4D(1836).

Shrub or tree 1^ m tall, erect, bushy, with branches erect or
ascending. Stems green to pale reddish brown, 4-lined when young,
soon 2-lined, eventually terete, internodes shorter than leaves; bark
becoming whitish then pale grey. Leaves sessile; lamina 20-70 x 5-
1 5 mm, narrowly elliptic to narrowly elliptic-oblong, the upper often
broader, plane, paler beneath with midrib prominent, not glaucous,
chartaceous, deciduous shortly before and during growth of new
shoots; apex acute to apiculate-obtuse or rarely rounded, base
narrowly cuneate to subangustate; venation: c. 8-1 2 laterals forming
looped intramarginal vein, sometimes with subsidiary laterals, ±
densely reticulate towards margins, tertiary venation dense and ±
obscure; laminar glands dense. Inflorescence up to c. 30-flowered
from up to 5 nodes, sometimes with flowering branches from up to
7 lower nodes immediately below or separated by sterile zone, the
whole broadly rounded-pyramidal to broadly cylindric; pedicels 4-
10 mm; bracteoles reduced foliar to triangular-subulate. Flowers
20-25(-37) mm in diam.; buds narrowly ovoid to narrowly ellip-
soid, acute to subacuminate. Sepals 3-4.5 x 1-2.2 mm, unequal,
varying from lanceolate, acute and basally united to oblong or
oblong-spathulate, rounded andr. 0.5 united, veins branched distally,
laminar glands basally linear, distally punctiform. Petals bright

135

yellow, not tinged red, 12-17 x 5-7 mm, c. 4 x sepals, oblanceolate-
unguiculate, cochleariform, rounded. Stamens 10-13 mm long, c.
0.7-0.8 x petals. Ovary 3-4 x 1 .5-2 mm, ellipsoid; styles 8-14 mm,
2.7-4.7 x ovary, basally separated and widely spreading-incurved.
Capsule (9-)10-12 x 7-8 mm, pyramidal-ovoid to ovoid-ellipsoid,
truncate to retuse, with horn-like persistent style bases, exceeding
sepals. Seeds yellowish brown, 1 .5-2 mm long; testa linear-reticu-
late to linear-foveolate (cf. Reynaud, 1991, f. 1, 3-6). 2n = 40
(Larsen, 1962; Borgen, 1969; Reynaud, 1986; Dalgaard, 1991).

Open rocky slopes, cliffs and ravines, disturbed ground, upper part
of litoral zone, relict Laurus forest; (20-)180-900(-1200) m.

Canary Islands (all western islands and possibly Fuerteventura),
Madeira. The only author to record its presence in Fuerteventura is
Voggenreiter (1974: 688, map). Naturalized in the Hawaiian Islands
(Maui) and southern California.

CANARY ISLANDS. Tenerife: San Diego de monte, 2 June 1855 (fl),
Bourgeau 1241 (JE, K); Orotava: The Quinta, St Ursula, March 1929 (fl),
Maude s.n. (BM); Montana* de Onaga, Barranco de las Huertas, 4 km above
San Andres, 8 April 1975 (fl), /, M. & P. Cannon 4669 (BM). Gran Canaria:
Barranco de los Tiles, April \%46(f\), Bourgeau 675 (BM, K); Miraflor, cerca
de Teror, 450 m, 14 April 1969 (fl), Kunkel 12747 (BM, H); El Brezal del
Palmital, near Moya, 21 July 1972 (fr), Melville & Bramwell 72/6 (K).
Gomera: below the Cumbre, 6 April 1861 (fl), Lowe Gl 10 (BM, K); Valle
Hermoso, May 1899 (fl), Murray s.n. (K). Hierro: Valverde, to the north, 12
February 1 858 (fl), Lowe H 1 1 8 (BM); W. of Frontera, 2 km E. of Sabinosa,
200 m, 12 April 1977 (fl), Jarvis & Murphy 260 (BM). La Palma: Barranco
del Rio, 400 m, 23 March 1905 (fl), Pitard 19 (FR, H, JE); near Los Llanos,
Barranco de los Angustias, 600 m, 20 April 1977 (fl), Jarvis, Gibby &
Humphries 397 (BM).

MADEIRA. Ribeiro de Joas Gomez, 300 & 800 m. May to September
1 865- 1 866 (fl), Mandon 36 (BM, JE, K); Santa, Ribeiro doTristEo gorge, N.
side, 150 m, 22 June 1985 (fl & fr), Press 1040 (BM).

HAWAIIAN ISLANDS (naturalized). Maui: East Maui, Kula, 960-
1080 m, 18 May 1985 (fl), Hobdy 2394 (BISH).

CALIFORNIA (naturalized). Sta Barbara Co., established at Montecito
and Santa Barbara (fide Munz, 1974: 519).

H. canariense is widely isolated from its nearest relatives in sect. 1.
Campylosporus on the African mainland, both morphologically and
geographically. Its inflorescence is most similar to that of H.
roeperianum, which occurs in West Africa and has densely reticulate
leaf venation; and for those reasons I at first regarded it as the nearest
relative of H. canariense (Robson, 1981: 68). Hagemann (1989:
242) has pointed out, however, that the growth form off/, canariense
is much nearer that of the mainly East African H. revolutum, which
also occurs in the Cameroon mountains and Fernando Poo. I agree
and now regard the Canary Island plant as most nearly related to the
broader-leaved form of H. revolutum in Ethiopia. H. canariense
differs from it inter alia by its broader leaves, more branched
inflorescence, smaller flowers with relatively small, pale-gland-
fringed sepals, narrower and not orange-tinged petals, fewer stamens
(the fascicles grouped 2+2-1- 1 ), trimerous ovary with relatively longer,
spreading and basally distinct styles, and relatively narrower capsule
with linear-foveolate rather than linear-reticulate seeds.

The type specimen of H. canariense has broad, rounded sepals,
whereas in that of H. floribun dum they are narrow and acute. Spach
(1836«, b), immediately followed by Webb & Berthelot (1836),
recognized an intermediate state; and, as was his wont, regarded the
difference between this group and the rest of Hypericum as worthy
of generic rank. Thus, in order of increasing sepalline acuity, Spach
and Webb & Berthelot, between them, described Webbia canariensis,
W. platysepala, W. heterophylla and W. floribunda. Later workers
found W. platysepala to be indistinguishable from W. canariensis,
and the intermediate 'species' W. heterophylla was also soon ignored;

136

N.K.B. ROBSON

Fig. 19 H. canariense: (a) habit; (b) leaf: (c) flower bud ('canariense'); (d) flower bud ('floribundum'); (e) sepal ('canariense'); (f) sepal ('floribundum');
(g) petal; (h) stamens and ovary; (i) anther; (j) capsule (a x 1/2; b x 1 ; c, d, h x 2; j x 3; e-g x 4; i x 6). All except d, f. Jarvis, Gibby & Humphries 397; d,
f. Cannon 4669.

STUDIES IN HYPERICUM

137

so H. canariense and H. floribundum assumed a rather spurious
distinctness. This view was encouraged by the geographical distri-
bution of the extremes: H. canariense in Tenerife, Gomera and
Hierro, H. floribundum in Gran Canaria, La Palma and Madeira.
Spach's intermediates do exist, however; and because of their exist-
ence and rather irregular distribution, it is not possible to recognize
more than one variable species, H. canariense L.

Sect. 22. ARTHROPHYLLUM Jaub. & Spach, ///.
pi orient. 1:44(1842).

Shrubs, low, compact and rounded to prostrate, up to r. 0.9 m tall,
eventually deciduous below but never leafless, glabrous, sometimes
with reddish to dark glands; branching lateral. Stems 2^(6)-lined
and ancipitous when young, soon 2-lined, not usually becoming
terete in first season, obscurely glandular; cortex greenish to reddish
brown; bark smooth, whitish grey. Leaves opposite, decussate,
sessile, free or perfoliate, deciduous at basal articulation; lamina
entire, with venation pinnate, closed, the tertiary densely reticulate;
laminar glands pale, punctiform; marginal glands dots pale, dense,
relatively large; ventral glands absent. Inflorescence 1-c. 40-flow-
ered, with branches dichasial/monochasial from 1-5 nodes, without
lower flowering branches; bracts and bracteoles reduced, entire or
black-gland-fringed. Flowers stellate, homostylous. Sepals 5, free
or basally connate, imbricate above, persistent, erect in fruit, with
margin entire or reddish- to black-gland-fringed; laminar glands
linear; marginal glands sessile or on short denticles or submarginal;
inframarginal glands absent. Petals 5, persistent, spreading and
twisting after flowering, with apiculus obsolete or absent; margin
entire; marginal glands absent; laminar glands linear to punctiform
or rarely absent. Stamen fascicles 3 (i.e. united 2+2+1), distinct,
persistent, with stamens 20-40; filaments basally united; anthers
yellow, gland amber; pollen types I, IV. Ovary with 3 axile placen-
tae, oo-ovulate; styles 3, free, bases separate; stigmas narrow or
narrowly capitate. Capsule 3-valved, subcoriaceous, with valves
longitudinally vittate. Seeds narrowly cylindric, ecarinate; testa
minutely rugulose.

BASIC CHROMOSOME NUMBER (x). Unknown.
HABITAT. Crevices in calcareous rocks, 100-1800 m.
DISTRIBUTION. S. Turkey, Syria, Lebanon.
5 species.

Key to sect. 22. Arthrophyllum

1 Sepals and bracts with marginal, reddish or black glands
Sepals and bracts without marginal glands

2(1)

Leaves free, lanceolate to narrowly ovate or oblong-elliptic, apex
rounded; sepals ovate to oblong, 0.35-0.5 united, marginal glands
black '• rupestre

Leaves perfoliate, almost completely united, broadly ovate to ob-
long-ovate, apex acute; sepals lanceolate, almost free or up to 0.35
united, marginal glands reddish 2. pamphylicum

3( 1 ) Leaves (at least middle and upper) with base cordate-amplexicaul;

flowers numerous, in dense corymbiform cymes

3. cardiophyllum

Leaves all with base cuneate to rounded; flowers 1-9, solitary or in
lax corymbiform cymes **

4(3) Sepals acute to shortly acuminate, lanceolate; leaves subcoriaceous.

venation not or scarcely prominent; flowers usually 2-9 (4. nanum)
5

Sepals rounded to subacute, oblong to ovate; leaves chartaceous,
venation prominent on both sides; flowers usually solitary
5. vacciniifolium

5(4) Stems erect, plant bushy; leaves ovate to broadly elliptic or orbicu-
lar; inner sepals triangular-lanceolate to narrowly oblong
4a. nanum var. nanum

Stems prostrate, plant appressed; leaves broadly to narrowly elliptic;
inner sepals ± broadly ovate 4b. nanum var. prostratum

1 . Hypericum rupestre Jaub. & Spach, ///. pi orient. 1: 44, tt. 21 ,
22 ( 1 842); Boiss., Fl. orient. 1 : 792 ( 1 867); R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 2nd ed. 21: 178 (1925); Stefanoff in
God. Agr.-les. Fak. Univ. Sofiya 11: 147 (1933), 12: 82 (1934), in
Pflanzenareale IV, 1: Karte 2a (1933); N. Robson in P. Davis, Fl
Turkey 2: 368, f. 11/5 (1967); Greuter, Burdet & Long, Med-
Checklist 3: 272 (1968). Types: Turkey [Mersin], 'in rupibus
abruptisCiliciae', [1834] (fl),/4wc7i6'r873 proparte (P-lectotype,
selected here; BM!, G!, K!); 'in rupibus montesTauri', 1 834 (fl),
Montbret s.n. (Fl, K!-syntypes). Jaubert & Spach described and
illustrated two varieties without attributing their cited specimens
to them. The Kew syntypes of both collections include twigs with
the upper leaves elliptic ('ovalifolia') and the lower ones
suborbicular ( 'rotundifolia').

Fig. 20D, Map 19.

H. rupestre [var.] a rotundifolium Jaub. & Spach, ///. pi. orient. 1:
44, t. 21 ( 1 842) ['rotundifolia'].Type (see above): Turkey, Cilicia,
1834 (fl), Montbret s.n. (Fl-holotype; K!).

H. rupestre [var.] p ovalifolium Jaub. & Spach, ///. pi. orient. 1: 44,
t. 22 (1842) ['ovalifolia}. Type (see above): Turkey, Cilicia,
[1834] (fl), Aucher 873 pro parte (P-holotype; BM!, G!, K!).

Icones: Jaub. & Spach, ///. pi. orient. 1: tt. 21, 22 (1842).

Shrub up to c. 0.3 m(?) tall, erect, bushy rounded, with branches

Map 19 Sect. 22. 1 . H. rupestre D; 2. H. pamphylicum A; 3. H.
cardiophyllum •; 5. H. vacciniifolium •.

138

N.K.B. ROBSON

erect or ± tortuous. Stems 4-lined and green when young; cortex
becoming red-brown and flattened in second year, then bark grey.
Leaves sessile; lamina 17^5 x 9-15 mm, lanceolate to elliptic or
suborbicular, paler beneath, midrib prominent proximally below, ±
glaucous beneath, rigidly coriaceous, deciduous during second year;
apex obtuse to rounded or (the lower) retuse, base broadly cuneate to
subattenuate; venation: c. 8-12 pairs of laterals, scarcely distinct
from tertiary reticulation. Inflorescence 9- 1 6-flowered from ( 1 )2(3)
nodes, subcorymbiform; pedicels 2.5-6 mm; bracteoles triangular-
subulate, margin black-glandular-ciliate. Flowers c. 20-25 mm in
diam.; buds elliptic, rounded. Sepals 2-3 x 0.7-1.2 mm, unequal to
subequal, 0.4-0.5 united, ovate to oblong-lanceolate or broadly
elliptic, subacute to rounded, margin distally or wholly with sessile
black glands; veins 5-7, subprominent. Petals bright yellow, not red-
tinged, 10-14 x c. 5-7 mm, c. 5 x sepals, obovate to oblanceolate,
asymmetrically retuse; laminar glands linear. Stamens 30-40, long-
est c. 9-13 mm, almost equalling petals. Ovary c. 3 x 1.5 mm,
ellipsoid; styles c. 10-12 mm long, 3^4 x ovary, widely curved-
ascending; stigmas narrowly capitate. Capsule c. 1 mm long,
exceeding sepals, narrowly ovoid. Seeds not seen.

Limestone cliffs; 150m.

Turkey (I9el).

TURKEY. I9el: Tarsus distr., gorge of Tarsus R. between Ulas and
Samlar. 150 m, 5 April 1957 (fl), Davis & Hedge D.26461 (BM, E, K).

Exsiccatae ofAucher 873 were labelled 'Syria' in error. H. rupestre
appears to be restricted to a small area in vilayet Icel, where it is rare.
Apart from the apomorphic black-glandular margins of the
bracteoles and sepals and the partial union of the sepals, H. rupestre
would seem to be the nearest species in sect. Arthrophyllum to H.
canariense morphologically and (except for 2. H. pamphylicum and
5. H. vacciniifolium) geographically. As H. canariense has sepals
with minute pale marginal glands, Spp. 1 and 2 ofsect.Arthrophyllum
may be regarded as having better developed marginal glands, whereas
Spp. 3-5 have apparently lost them.

2. Hypericum pamphylicum N. Robson & P. Davis in Notes R.
hot. Gdn Edinb. 38: 1 04 ( 1 980); N. Robson in P. Davis, Fl. Turkey
10: 96 (1988); Greuter, Burdet & Long, Med-Checklist 3: 270
(1986). Type: Turkey, Antalya, Alarahan, 7 km inland from coast
road between Manavgat and Alanya, beneath castle, 100 m, 11
May 1979 (fl), Matthew, Baytop & Siitlupinar 9599 (BM!-
holotype; E!, ISTF, K!-isotypes).

Fig. 20B, Map 19.

Shrub c. 0.14-0.25 m tall, with branches decumbent. Stems slightly
2-lined and green when young, soon terete; cortex becoming reddish
brown, then bark grey in second year. Leaves in almost completely
perfoliate pairs; lamina 18-35 x 14-35 mm, broadly ovate or
broadly oblong-ovate, paler beneath, midrib slightly prominent
beneath, glaucous, subcoriaceous, deciduous during second year;

apex acute to apiculate-obtuse or rounded, base united; venation: 3-
4 pairs of laterals, scarcely distinct from tertiary reticulation.
Inflorescence 5-20-flowered from (1)3^4 nodes, hemispheric to
subcorymbiform; pedicels 3-4 mm; bracteoles lanceolate-subulate,
margin red-glandular-denticulate. Flowers c. 20 mm in diam.; buds
elliptic, obtuse to subacute. Sepals 3-3.5 x 0.7-1 mm, unequal,
almost free to 0.35 united, lanceolate to narrowly oblong, obtuse to
rounded, margin red-glandular-denticulate, veins 5, not or scarcely
prominent. Petals bright yellow, not red-tinged, 1 1-12(-14) x 3 mm,
c. 4 x sepals, lanceolate to narrowly oblong, rounded; laminar
glands striiform. Stamens 21-26, longest 10-14 mm long, about
equalling petals. Ovary 3x1.5 mm, narrowly ovoid; styles c. 1 0 mm
long, c. 3.5 x ovary, narrowly curved-ascending; stigmas narrowly
capitate. Capsule and seeds unknown.

Limestone rocks; 100 m.

Turkey (Antalya).

TURKEY. Antalya: Alarahan, 7 km inland from coast road between
Manavgat and Alanya, 100 m, 11 May 1979 (fl), Matthew, Baytop &
Sutlupinar 9599 (BM, E, ISTF*, K).

H. pamphylicum has been collected only once. It is clearly closely
related to H. rupestre, in relation to which all its characters are
apomorphic with the possible exception of the red (as opposed to
black) glands.

3. Hypericum cardiophyllum Boiss., Fl. orient. 1: 791 (1867); R.
Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 178
(1925); Boul., Fl. Liban: 68 (1930); Stefanoff in God. Agr.-les.
Fak. Univ. Sofiya 11: 147 (1933), 12: 82 (1934), \nPflanzenareale
IV, 1: 2a (1933); J. Thiebaut, Fl. Lib.-Syr. 1: 139 (1936); N.
Robson in P. Davis, Fl. Turkey 2: 367, f. 1 1/4 (1967); Mouterde,
Nouv. Fl. Liban. Syrie 2: 521 (1970); Greuter, Burdet & Long,
Med-Checklist 3: 265 ( 1 986). Types: Syria, 'Darkusch ad Orontem
in via ab Antiochia ad Aleppo', June 1846?, Boissier s.n. (G!-
syntype); Turkey, Gaziantep, 'Assy prope Aintab, etiam prope
Urfa', 600 m, 24 June 1865 (fl & fr), Haussknecht s.n. (G!-
lectotype, selected here; K!-isolectotype). Haussknecht collected
this species in two localities, but it is not clear from which the
type came; perhaps specimens from both localities were mounted.
He subsequently distributed exsiccatae of both collections: 'Ad
rupes Assy prope Aintab' (Haussknecht 603) and 'In cacumine
Montis Nar Facub, prope Orfa' (Haussknecht 663). These may be
duplicates of the lectotype.

Map 19.

Shrub 0. 12-0.9 m tall, erect, bushy, rounded, with branches erect to
ascending. Stems 2-lined and green when young; cortex loosening
and becoming whitish in second year, then bark pale grey. Leaves
sessile; lamina (15-)20-45 x 10-30 mm, oblong or elliptic to ovate
or lanceolate, almost concolorous, midrib scarcely prominent, ±

STUDIES IN HYPER/CUM

139

Fig. 20 A. H. cardiophyllum: (a) habit; (b) sepal; (c) petal; (d) capsule. B. H. pamphylicum: (e) habit; (0 sepal. C. H. vacciniifolium: (g) habit; (h) sepal.
D. H. rupestre: (i) habit; (j) sepal. E. H. nanum: (k) habit; (1) sepal (a, e, g, k x Vr, all others x 4). A. Post s.n. B. Mathew et al. 9599. C. Siehe 226. D.
Davis & Hedge 26461. E. Norris s.n.

140

N.K.B. ROBSON

glaucous especially beneath, thinly coriaceous, deciduous before (?)
growth of new shoots; apex obtuse to rounded, base truncate to
cordate-amplexicaul; venation: 5-6 pairs of laterals, scarcely dis-
tinct from tertiary reticulation. Inflorescence c. 1 0-40-flowered
from 3(-5) nodes, corymbiform to broadly hemispherical; pedicels
2-3 mm; bracteoles triangular-subulate, margin entire. Flowers c.
16-20 mm in diam.; buds elliptic, obtuse. Sepals 2-4 x 0.9-2 mm,
subequal, free, triangular-ovate to oblong-lanceolate, acute to suba-
cute or sometimes rounded, entire, glands submarginal; veins 5, not
prominent. Petals bright yellow, not tinged red, 9-12 x (2.5-)5-7
mm, 3-4.5 x sepals, obovate to elliptic, rounded; laminar glands
striiform to punctiform. Stamens 20-30, longest 9-12 mm, about
equalling petals. Ovary c. 1.5-2 x 1-1.5 mm, ovoid; styles c. 9-1 1
mm long, c. 4 x ovary, widely spreading; stigmas narrow. Capsule
4.5 x 2.5-3.5 mm, very narrowly ovoid to cylindric, truncate, with
persistent horn-like style bases, exceeding sepals. Seeds yellowish
brown, c. 1 .2 mm long.

Open calcareous rocks; 500-1000 m.

TURKEY. Gaziantep: Osmaniye to Gaziantep, 48 km W. of Gaziantep,
950- 1000m, 24 June 1953 (fl), Huber-Morath 12087 (BASBG); c. 4km N.
of Halfeti, 500 m, 24 May 1983 (fl), Sorger%3-5-3> (W);Aintab [Gaziantep],
ad rupes Assy prope Aintab, 24 June 1 865 (fl), Haussknecht 603 (BM, E, G,
JE, K); Aintab, 15 June 1882 (fr), Post s.n. (BM). Urfa: 5 km NE of Halfeti,
650m, 30April 1980 (e.ft),Sorger 80-14-48 (W);Orfa [Urfa], in rupestribus
calcareis, 24 June 1867 (fl), Haussknecht s.n. (BM); in cacumine mentis Nar
Facub, prope Orfa, 15 May 1865 (fl), Haussknecht 663 (JE).

SYRIA. Latakia |A1 Ladhiqiyah]: ad rupes prope Latakieh, June 1846
(fl), Boissier (K, UPS); no precise locality, 1 846 (fl), Pinard s.n. (BM, H, K);
Darkhush [on R. Orontes], June 1865? (fl), Boissier s.n. (G).

H. cardiophyllum has most of the characters of H. canariense in
miniature or in reduced numbers, except for the corymbiform inflo-
rescence and differently shaped glaucous leaves. The truncate to
cordate leaf-base and entire acute sepals distinguish it from other
species in sect. Arthrophyllum. The flowers are somewhat smaller
and more crowded than those of 1 . H. rupestre, and it lacks the dark
glands found in that species.

4. Hypericum nanum Poir., Encycl., Suppl.3: 699 (1814); Choisy,
Prodr. monogr. Hyperic. : 49 ( 1 82 1 ), in DC., Prodr. 1: 549 ( 1 824);
Spach in Annls Sci. nat. (Bot.) II, 5: 357 (1836); Jaub. & Spach,
///. pi. orient. 1: 46, t. 23 (1842); Boiss.,F/. Orient. 1: 792 (1867);
R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 178
(1925); Boul., Fl. Liban: 69, t. 68 f. 3 (1930); Post, Fl. Syria 2nd
ed. 1: 229 (1932); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya
10: tt. 1 f. 4, 2 f. 8, 3 f.10 (1932), 11: 146 (1933), 12: 82 (1934),
in Pflanzenareale IV, 1 : Karte 2a ( 1 933); J. Thiebaut, Fl. Lib. -Syr.
1: 140 (1936); Oppenheimer & Evenari in Bull. Soc. hot. Geneve
31: 324 (1940); Zohary, Fl. Palaestina 1: 222 (1966); Mouterde,
Nouv. Fl. Liban Syrie 2: 521, t. 224 f. 4 (1970); Greuter, Burdet
& Long, Med-Checklist 3: 269 ( 1 986); N. Robson in Cullen et al.,
Eur. GdnFl. 4: 70, ff. 1 1.3, 1 1.13 (1995).Type: Lebanon, 'Syria',
no precise locality [Cossaye?], de Labillardiere (P-holotype; ?H!
- see below).

Fig. 20E, Map 20.

Icon: Jaub. & Spach, ///. pi. orient. 1: t. 23 (1842) (var. nanum).

Map 20 Sect. 22. 4. H. nanum • specimens, D records.

Shrub c. 0.15-0.3 m tall, erect, bushy, rounded ('hemispherical'),
with branches erect or ± tortuous or sometimes prostrate with stems
appressed against rocks. Stems 4-lined and green when young, soon
2-lined to terete; bark greyish to whitish grey. Leaves sessile; lamina
5-20(-25) x 4-15(-18) mm, ovate or broadly oblong-ovate or
orbicular to broadly or narrowly elliptic, somewhat paler beneath,
midrib prominent beneath, ± glaucous when young, sometimes
persistently so beneath, subcoriaceous, deciduous during second
year; apex obtuse or subapiculate to rounded or rarely retuse, base
rounded to cuneate or more rarely shortly angustate; venation: 4-6
pairs of laterals, scarcely distinct from tertiary reticulation. Inflores-
cence 2-9-flowered from 1-2 nodes, rounded-corymbiform; pedicels
4.5-9 mm; bracteoles triangular-subulate, margin entire. Flowers c.
12-20 mm in diam.; buds ellipsoid, rounded. Sepals 2-3(-3.5) x
0.5-1 mm, subequal to unequal, free or up to 0.2 united, triangular-
lanceolate to narrowly oblong or rarely ovate, acute or subacuminate
to subacute, margin entire or with distal marginal glands ± protrud-
ing, veins 5, at least midrib prominent. Petals bright yellow,
(6-)8-10(-12) x c. 2-4.5 mm, c. 3^4- x sepals, oblanceolate to
narrowly cuneate, asymmetrically retuse; laminar glands striiform
to punctiform or absent. Stamens 30-40, longest c. 8-10 mm, about
equalling petals. Ovary c. 2-3 x 1-1.5 mm, narrowly ovoid; styles
9-12 mm long, c. 3-5 x ovary, narrowly curved-ascending; stigmas
narrow. Capsule 5-6.5(-8) x 3^.5(-6) mm, cylindric-ovoid, apically
impressed. Seeds not seen.

Calcareous rocks; (750)900-1800 m.

Lebanon (Lebanon and Antilebanon mts), Syria (Antilebanon).

H. nanum exists in two growth forms which, unless or until they are
shown to merge, are best treated as varieties.

4a. Hypericum nanum var. nanum

Stems erect, forming rounded bushes. Leaves c. 10-20 mm long,
ovate or oblong-ovate or orbicular to broadly elliptic, subapiculate
or rounded to retuse. Inflorescence 3-9-flowered. Sepals (2-)2.5-
3.5 mm long, the inner triangular-lanceolate to narrowly oblong.

Range of species; 750-1800 m.

STUDIES IN HYPER1CUM

LEBANON. Qadisha gorge below Becharre [Bsharri|, 900-1050 m, 19
August 1945 (fr),Davis 10149 (BM,K);Cossaye [Kassayer?], ( 1787] (fl),de
Labillardiere s.n. (H)-cf. type. Antilebanon: Wadi Jemayli (above Baalbek),
1350-1800 m, 23 June 1943 (fl), Davis 6582 (BM, K); near Rukhby
[Rukhbi], 14 July 1890 (fl), Post s.n. (E, K).

SYRIA. Antilebanon circa Zebdaine [Zebdani | prope Damascum, decus
parientum imminentium vallis Uod e Uom, 1 200 m, 7 June 1 855 (fl), Kotschv
68 (BASBG, BM, K, UPS); Hermon, 1200 m, May 1945? (fl), Norris (BM).

The Bsharri collection has larger flowers and leaves than the others,
the leaves having a distinct angustate base and (mostly) retuse apex.
The collection was originally determined as H. rupestre, and the
leaves do resemble those in Jaubert & Spach's ( 1 842) figure of their
var. ' ' rotundifolia '; but the sepals and bracteoles are entire as in H.

4b. Hypericum nanum var. prostratum Boiss., Fl. orient. 1: 792
(1867); Post, Fl. Syria 2nd ed. 1: 229 (1932); Rech. f. in Ark.
Bot. 5: 292 (1960); Mouterde, Nouv. Fl. Liban Syrie 2: 521
( 1970). Type: Lebanon, Antilebanon, rochers au bordsdu Barrada
entre Bessime [Bassima] et Ain Fige [Ain el Fiji], 18 May 1817
(fl), Gaillardot 1676 (G-holotype; JE!).

Stems prostrate, appressed to rocks. Leaves 5-12 mm long, broadly
to ± narrowly elliptic, obtuse to rounded. Inflorescence 2-5-flow-
ered. Sepals 2-2.5 mm long, the inner ± broadly ovate.

LEBANON. Antilebanon: prope A'm Yunun, 1600 m, 21 May 1910 (fl),
J.&F. Bornmiiller 11519 (BM, JE); Falita (above Nabh), 1650 m, 21 June
1943 (fl), Davis 6582 (BM, K).

SYRIA. Antilebanon: S. of Ain el Fiji, 20 km NW of Damascus along
Barada valley, 5 May 1963 (fl), Barkoudah 741 (E).

The shape and angustate base of the leaves of the Bsharri population
(q.v. supra) suggest that H. nanum is directly related to H. rupestre
rather than to H. cardiophyllum, in which the entire sepals, smaller
flowers and cordate leaf-bases may therefore be regarded as
apomorphic in relation to H. rupestre.

5. Hypericum vacciniifolium Hayek & Siehe in Annln naturh.
Mas. Wien 28: 159, t. 1 1 f. 3 (1914); Stefanoff in God. Agr.-les.
Fak. Univ. Sofiya 11: 146(1933), 12: 82(1934), \nPflanzenareale
IV, 1: Karte 2a (1933); N. Robson in P. Davis, Fl. Turkey 2: 368
(1967); Greuter, Burdet & Long, Med-Checklist 3: 274 (1986);
N. Robson in Cullen et al., Eur. Gdn FL 4: 70 (1995). Type:
Turkey, Vil. Adana [I?el], Sandjak Mersina, bei Efrenk, 1500 m,
Junel912(fl),5/e/i£'226(W-holotype;BM!,E!,JE!,Z!-isotypes).

Fig. 20C, Map 19.

H. nanum var. uniflorum Bornm. in sched.

Icon: Hayek & Siehe in Annln naturh. Mus. Wien 28: 159, t. 1 1 f. 3
(1914).

jo

Shrub 0.08-0.2 m tall, erect, bushy, rounded, with branches ±
tortuous. Stems 2-lined when young, soon terete; bark greyish brown
to whitish grey. Leaves sessile or with pseudopetiole up to c. 0.7 mm;
lamina 6-15 x 3.5-9 mm, elliptic or oblong-elliptic to obovate,
somewhat paler but not or scarcely glaucous beneath, midrib and
reticulate venation ± prominent on both sides, chartaceous, decidu-
ous during second year; apex obtuse or subapiculate to rounded,

141

base cuneate to angustate or shortly pseudopetiolate; venation: 3-6
pairs of major and minor laterals, ± distinct from tertiary reticula-
tion. Inflorescence l-3(-9)-flowered, from 1-2 nodes,
rounded-corymbiform when several-flowered; pedicels 4-7 mm;
bracteoles triangular-subulate, margin entire. Flowers c. 15-18 mm
in diam.; buds ellipsoid, rounded. Sepals 2-4 x 1.3-1.7 mm, un-
equal, shortly united, oblong to ovate, subacute to rounded, margin
entire, glands submarginal, veins 3-5, not prominent. Petals bright?
yellow, 10-12 x 4-5 mm, 3-4 x sepals, oblong-lanceolate, un-
equally retuse; laminar glands linear to punctiform. Stamens c. 20,
longest c. 11-1 2 mm, about equalling petals. Ovary c. 2.5 x 1.5mm,
narrowly ovoid-ellipsoid; styles 8-9 mm long, c. 3-4 x ovary,
narrowly curved-ascending; stigmas narrow. Capsule (immature)
ovoid. Seeds not seen.

Limestone cliffs; 1000-1500 m.

Turkey (E. Cilicia).

TURKEY. I9el: Mut-Kirobasi,33km, 1260m, 14 June 1 950 (fl), Attila
in Hub.-Mor. 1 1489 (BASBG); Mut to Biiyiik Egri Dagh, 1500 m, 12 May
1965(fl),C00^<ft70ws883(E,K);Lamas-Schlucht, 1000m, 12 June 1912
(fl), Siehe s.n. (JE).

H. vacciniifolium is closely related to the Libano-Syrian H. nanum,
differing essentially from it in the thinner leaves with prominent
venation and angustate to pseudopetiolate base and the smaller
number of stamens, and usually in the fewer (often only 1 )-flowered
inflorescence and relatively broader, less acute sepals. It is confined
to a small area of the Taurus Mountains in eastern Cilicia (vilayet
Icel).

Sect. 23. TRIADENIOIDES Jaub. & Spach, ///. pi.
orient. 1:49(1842).

Shrubs or shrublets, erect to prostrate, up to c. 0.6 m tall, deciduous
below but never wholly leafless, glabrous or rarely with innovations
and lower surface of leaves puberulous, sometimes with dark glands;
branching lateral. Stems 4-6-lined and ± compressed (ancipitous)
when young, becoming terete in second season, eglandular or rarely
with a few dark glands; cortex green to red; bark smooth to finely
ribbed-striate, grey-brown to red-browa Leaves opposite or 3-
whorled, decussate, sessile or petiolate, free, deciduous at basal
articulation; lamina entire, with venation pinnate and partly open or
1 -nerved, the tertiary reticulation obscure (or absent?); laminar
glands pale or rarely dark, punctiform; submarginal dark glands
rarely present; marginal or intramarginal gland dots pale, dense or
rather sparse, similar to laminar ones; ventral glands absent. Inflo-
rescence 1-13-flowered, with branches dichasial/monochasial
(sometimes subopposite) from 1-2 nodes or subumbellate, without
lower flowering branches; bracts and bracteoles reduced or bracts
foliar, entire. Flowers stellate, homostylous.5f/?a/.v 5, free or slightly
basally connate, persistent, erect to recurved in fruit, with margin
entire or rarely dark-gland-fringed; veins 3-7; laminar glands pale,
linear to punctiform; marginal glands on denticles or subsessile or
absent; submarginal glands absent or dark; inframarginal glands
absent. Petals 5, persistent, spreading or erect but not twisting after
flowering, with apiculus subterminal, short or absent; margin entire;
marginal glands absent, laminar glands linear to punctiform, pale or
rarely reddish to black. Stamen fascicles 3 (i.e. united 2+2+1),
distinct, persistent, with stamens (10-)20-40(-60); filaments ba-
sally or up to c. 0.4 united; anthers yellow, gland amber or rarely
black; pollen types IV, VI. Ovary with 3 loosely axile placentae, each
many- to few-ovulate; styles free, bases distinct, contiguous; stig-
mas narrowly to scarcely capitate. Capsule 3-valved, coriaceous to
subcoriaceous, with valves ± prominently longitudinally vitiate or

142

verrucose. Seeds narrowly cylindric, narrowly or not carinate, proxi-
mally ± complanate; testa reticulate-foveolate to linear-foveolate or
rugulose.

BASIC CHROMOSOME NUMBER (X). 8; ploidy 2.

HABITAT. Limestone or less commonly conglomerate or granitic
rocks or slopes; 0-1900 m.

DISTRIBUTION. Socotra, south Turkey, Syria, Lebanon.
5 species.

Key to sect. 23. Triadenioides

1 Leaves discolorous, paired; dark glands absent 2

Leaves concolorous, 3-whorled or with dark glands 4

2(1) Leaves whitish-puberulous beneath and young parts fawn-
puberulous; all leaves petiolate, triangular-ovate to oblong-ovate
1. fieriense

Leaves pruinose to whitish-puberulous beneath or whole plant
glabrous; all or some leaves sessile, elliptic or oblong to obovate
3

3(2) Flowers 1 (2), terminal and in axils of older leaves; sepals ensiform;
capsule valves vitiate; habit erect 2. scopulorum

Flowers 7-13 in terminal subumbellate inflorescence: sepals elliptic
or oblong-oblanceolate to oblong; capsule valves verrucose; habit
spreading or straggling 3. tortuosum

4( 1 ) Leaves in whorls of 3, without dark glands; usually dense ± rounded
shrub 4. ternatum

Leaves paired, with inframarginal dark glands; straggling to pros-
trate shrublet 5. pallens

1 . Hypericum fieriense N. Robson in Bull. nat. Hist. Mus. Land.
(Hot.) 23: 68 (1993). Type: Socotra, Hagghiher Mountains
(12°35'N, 54°03'E), below Fieri peaks, 1350 m, 21 April 1967,
Smith & Lavranos 475 (K!-holotype & isotype).

Fig. 21E, Map 21.

Shrub c. 1 m tall, much branched, flat-topped, with branches ascend-
ing, dark glands absent. Stems fawn-puberulous, soon regularly

N.K.B. ROBSON

4-lined; cortex green?; bark finely ribbed-striate. Leaves opposite,
the pairs initially united at base, petiolate, with petiole 4-6 mm long,
densely to rather sparsely fawn-puberulous with scattered stellate
hairs; lamina 10-17 x 7-12 mm, triangular-ovate to oblong-ovate,
glabrous above, paler and densely fawn-puberulous beneath with
scattered, often tufted villous hairs, midrib and laterals prominent
beneath and ± impressed above, coriaceous; apex subobtuse to
rounded, margin inrolled, base broadly cuneate to shortly angustate
or truncate; venation: 4-5 pairs of laterals, closed or lower 1 -2 pairs
free, without or with 1-3 cross-veins distally; laminar glands dense,
prominent on upper surface, intramarginal glands dense. Inflores-
cence 3-5-flowered, from 1-2 distant or ± close nodes, the
inflorescence thus more or less subumbellate; pedicels 8-9 mm,
bracts and bracteoles linear. Flowers 9-10 mm in diam.; buds
narrowly ovoid, acute. Sepals in fruit recurved, 4-5 x c. 1.5 mm,
linear-lanceolate, acute, margin entire, coriaceous; veinsc. 7, slightly
prominent; laminar glands linear, all pale. Petals incomplete in
specimen seen. Stamens c. 60?, with filaments (in each fascicle)
united above base. Ovary not seen. Capsule 6-7.5 x 4.5-6 mm,
pyramidal-ovoid, truncate, with persistent divergent horn-like style
bases, coriaceous, with valves finely longitudinally vitiate, exceed-
ing sepals. Seeds dark brown, c. 1.1 mm long (immature?); testa
foveolate-reticulate.

'Low scrub among Dracaena cinnabari trees'; 1350 m.

Socotra (Hagghiher Mts).

SOCOTRA. Hagghiher Mts, 12°35'N,54°03'E, below Fieri peaks, 1350
m, 21 April 1967 (fr), Smith & Lavranos 475 (K).

This strikingly distinct species is known from only the above
collection. It is clearly related to the other Socotran low shrubs, H.
scopulorum and//, tortuosum, bul differs from theminteralia in leaf
shape and inflorescence form and, particularly, in the presence of an

Map 21 Sect. 23: 1. H. fieriense •; 2. H. scopulorum O; 3. H. tortuosum •.

STUDIES IN HYPERICUM

indumentum on young vegetative parts and the lower surface of the
leaves.

2. Hypericum scopulorum Balf. f. in Proc. R. Soc. Edinb. 11: 502
(1882), in Trans. R. Soc. Edinb. 31: 27, t. 4A (1888), in Forbes,
Nat. Hist. Socotra: 456 (1903); Vierh. in Denkschr. Akad. Wiss.
Wien 71: 389 (1907); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 178 (1925) ['scopulosum']; N. Robson
in Kew Bull. 12: 434, 444 (1958); Moggi & Pisacchi in Webbia
22: 268, f. 10, carta 6 (1967). Type: Socotra, Passhohe oberhalb
Kischen, 1000 m, 6 May 1881, Schweinfurth 756 (E-lectotype,
Moggi & Pisacchi, 1967); iiber Kischen, 900 m, 1 May 1881,
Schweinfurth 622 (E, LE-syntypes); in montibus Haggier [Debra
hi-hon], 900 m, February-March 1880 (fr), I.B. Balfour [with
Cockburn & Scott] 405 (BM!, E-syntypes).

Fig. 21A, Map 21.

Icon: Moggi & Pisacchi in Webbia 22: 270, f. 10 (1967).

Shrub 0.3-c. 1 .3 m tall, much branched, flat-topped, with branches
erect to ascending; wholly glabrous or (especially upper) pruinose to
whitish-puberulous beneath; dark glands absent. Stems persistently
4(6)-lined; cortex reddish?; bark finely ribbed. Leaves opposite,
free, sessile or rarely petiolate with petiole up to 2 mm long; lamina
10-27 x 6-12(-17) mm, elliptic to oblong, paler beneath, ± densely
glaucous on both sides, coriaceous; apex subapiculate-obtuse to
rounded, margin plane to subincrassate, base narrowly cuneate to
subangustate; venation: 4-6 pairs of laterals, closed or lower 1-2
pairs free, without cross-veins; laminar glands ± dense, not or
slightly prominent on both sides, intramarginal glands dense. Inflo-
rescence 1 (2)-flowered, terminal and from axils of older leaves of
current shoot; bracts foliar, bracteoles lanceolate to linear-subulate,
deciduous; pedicels 3-8 mm long, slender. Flowers 10-14 mm in
diam.; buds narrowly ellipsoid, acute. Sepals5-6.5 x 1-1 .3 mm, free
or very shortly connate, distally outcurving in fruit, ensiform, acute,
coriaceous, margin entire; veins 7, slightly prominent; laminar
glands punctiform. Petals golden yellow to orange yellow (old?), not
tinged red, 6-7 x 2-3 mm, 1.1-1.2 x sepals, oblong-oblanceolate,
rounded. Stamens c. 30, longest c. 3.5 mm, c. 0.5 x petals. Ovary c.
1.3 x 0.6 mm, narrowly ellipsoid, obtuse; styles 2-3.5 mm long, 2-
3 x ovary, curved-ascending. Capsule c. 4 x 3.5 mm, broadly
ellipsoid, obtuse, subcoriaceous (enclosed by old, distally twisted
petals), with valves longitudinally vittate, shorter than sepals. Seeds
dark brown, 1.3-1.6 mm long; testa foveolate-reticulate.

High-altitude thicket on rocky or grassy slopes; 775-1322 m.

Socotra (Hagghiher Mts).

SOCOTRA. Adho Demalu, 900 m, 16 March 1953 (fl), Popov GP/SO/
262 (BM, EA); Jebel Shihali, 20 April 1967 (fl & fr), Smith & Lavranos 446
(BM, K); Debra hi-hon, 900 m, 25 August 1956 (fl), Gwynne 138 (BM);
Aduno pass, 775 m, 6 March 1989 (fl), Miller et al. M.8671 (E, K).

Of the three endemic Socotran Triadenioid species, H. scopulorum
has the most primitive leaves (glabrous and mostly sessile); but it is
less woody than H.fierense, and the inflorescence and fruit are more
reduced. According to the label of Smith & Lavranos 446, it is
associated with Croton eleagnoides Balf. f.

143

3. Hypericum tortuosum Balf. f. in Proc. R. Soc. Edinb. 11: 502
(1882), in Trans. R. Soc. Edinb. 31: 28, t. 4B (1888), in Forbes,
Nat. Hist. Socotra: 457 (1903); Vierh. in Denkschr. Akad. Wiss.
Wien 71: 389 (1907); G.B. Popov in / Linn. Soc. (Bot.) 55: 714
(1957); N. Robson in Kew Bull. 12: 434, 444 (1958); Moggi &
Pisacchi in Webbia 22: 269, f. 1 1, carta 6 (1967). Type: Socotra,
Passhohe iiber Kischen, 1000 m, 6 May 1 88 1 , Schweinfurth 757
(E-lectotype, Moggi & Pisacchi, 1967); no precise locality ['With
the foregoing species (H. scopulorum) on the Haghier range at a
high elevation' (600 m)], February-March 1880, I.B. Balfour
[with Cockburn & Scott] 607 (E-syntype).

Fig. 21B, Map 21.

Icon: Moggi & Pisacchi in Webbia 22: 271, f. 1 1 (1967).

Shrub or shrublet toe. 0.5 m tall, much branched, ± round-topped or
trailing, with branches divergent-ascending, flexuous; wholly gla-
brous or with young stems and leaves beneath densely and minutely
papillose; dark glands absent. Stems persistently 4-lined; cortex
reddish; bark finely longitudinally ribbed. Leaves opposite, free, all
sessile or lower petiolate, with petiole up to 4 mm long; lamina 8-
1 7(-23) x 4-1 1 (- 1 5) mm, obovate or oblanceolate to elliptic-oblong
or suborbicular, paler beneath, glaucous on both sides (densely so
beneath), coriaceous; apex obtuse (or subacute?) to rounded, margin
revolute, base cuneate; venation: 4-5(-8) pairs of laterals, closed,
without cross-veins but sometimes with rather obscure tertiary
reticulation; laminar glands dense, obscure, not prominent or slightly
so beneath or on both sides; intramarginal glands rather dense,
obscure. Inflorescence 1-c. 35-flowered, terminal, subumbellate,
occasionally with single axillary flower from lower node; upper-
most leaf pair modified as bracts, oblong, amplexicaul; bracteoles
minute, basal; pedicels 4-8 mm long, slender. Flowers c. 10 mm in
diam.; buds ellipsoid, obtuse. Sepals 4-6 x (1-)1. 5-2.5 mm, free,
imbricate, unequal, elliptic to oblong-oblanceolate or oblong, acute
to rounded, thinly chartaceous, margin entire; veins 7(-l 1), promi-
nent, pinnately branched; laminar glands absent. Petals bright yellow,
not tinged red, 6-7 x 2-3 mm, 1 .5-2 x sepals, elliptic to oblanceolate,
rounded. Stamens c. 25, longest 4-7(-7) mm, c. 0.7 x petals. Ovary
1.5-2 x 1-1.7 mm, ovoid-ellipsoid, acute; styles 3-5 mm long, 2-
2.5 x ovary, curved-ascending. Capsule c. 5 x 2.5 mm, ellipsoid,
acute, subcoriaceous, with valves glandular-verrucose (vesiculate),
about equalling sepals. Seeds dark brown, c. 1.2 mm long; testa
rugulose.

Among rocks and crags (sometimes limestone) on hillsides; (33-)
200- 1200m.

Socotra (Hagghiher Mts, Reighid, extreme west).

SOCOTRA. Reiged [Reighid], 750 m, 1 2 March 1953 (fl & e. fr), Popov
GP/So/213 (BM, EA); Jebel Rhugid, 600 m, 8 February 1990 (fl). Miller et
al. M. 10340 (E, K*, UPS*); Shihali, c. 9 km SSE of Hadiboh, 1 150 m, 6
March 1989 (fl & fr),A//7fcret al. M.8680 (E); Ras Shoalo, 33 m, 19 February
1953 (fl), Popov GP/So/143 (BM); 6 km E. of Ras Kattanahan, 200 m, 17
January 1994 (fl & e. fr), Thulin & Gifri 8579 (K, UPS); 16 km SE of
Qalansiyeh, 700 m, 24 January 1994 (fl), Thulin & Gifri 8723 (K, UPS); no
precise locality, 1897 (fl), Mr & Mrs T. Bent s.n. (K). Also recorded from
Adho Dimellus in Forbes (1903) and by Vierhapper (1907) and from near the

144

N.K.B. ROBSON

Fig. 21 A. H. scopulorum: (a) habit; (b) leaf: (c) sepal; (d) petal; (e) capsule. B. H. tortuosum: (f) habit. C. H. ternatum: (g) habit; (h) leaf; (i) anther. D.
H. pollens: (j) habit; (k) leaf; (1) sepal; (m) petal; (n) capsule. E. H.fierienxe: (o) habit; (p) leaf; (q) sepal; (r) capsule (a, f, g, j, o, x '/z; b, p x 2; h, k, x 3;
c-e, 1-n, q, r x 5; i x 10). A. Smith & Lavranos 446. B. Popov 213. C. Davis 15514. D. Davis & Polunin 25968. E. Smith & Lavranos 475.

STUDIES IN HYPER1CUM

summit of Jebel Serai (1322 m) and Adho Pass (877 m) by Simony, 18
February 1 899 (fide Vierhapper, 1907).

H. tortuosum is apomorphic relative to the other two Socotran
species of sect. Triadenioides in its less woody, spreading to pendu-
lous habit, smaller size, larger sepals and verrucose capsules; and its
subumbellate inflorescence and usually petiolate leaves are also
specialized relative to H. scopulorum. The population originally
described (from the central mountains) has leaves (except the bracts)
all, or at least the lower, petiolate with 4-5 lateral veins and no
visible tertiary reticulate venation, and the young stems and lower
leaf surface are smooth to undulate; whereas the western lowland
population has wholly sessile, larger leaves and sometimes larger
flowers, the leaves may have up to 8 pairs of lateral veins and
obscurely visible tertiary reticulate venation, and the young stem
and the leaves beneath are sometimes minutely papillose.

The western population therefore approaches H. scopulorum in
several characters; but in others (e.g. the prostrate habit) it is typical
of H. tortuosum. It would seem, then, to be the remains of the stock
from which the above two species diverged, an interpretation that is
supported by the presence in some leaves of more numerous lateral
veins than is usual in either species, thus showing a tendency
towards H. socotranum (sect. 1 . Campy losporus). These characters
of the western population do not, however, provide a clear-cut
separation of the two populations, e.g. the lower leaves of the central
mountain population may be shortly petiolate (Miller et al. M.8680),
or leaves at several upper nodes may be wholly sessile (Balfour,
1 888: t. 4B). I therefore hesitate to treat these populations as separate
taxa, preferring to regard them as geographically distinct, respec-
tively plesiomorphic and apomorphic parts of an almost continuous
morphocline.

4. Hypericum ternatum Poulter in Notes R. bot. Gdn Edinb. 21:
181 (1954); N. Robson in P. Davis, Fl Turkey 2: 368, f. 11/6
( 1 967); Greuter, Burdet & Long, Med-Checklist 3: 273 ( 1 986); I.
Hagemann in Flora 183: 288, ff. 60-63 (1989). Type: Turkey,
Antalya, distr. Gebiz, Bozburun Dag nearTozlu Cukur yayla, 24
July 1 949 (fl), Davis & BilgerD. 1 5580 (K!-holotype; E!-isotype).

Fig. 2 1C, Map 23.

Icon: I. Hagemann in Flora 183: 288, f. 60 (1989).

ShrubletQM-Q. 1 5(-0.27) m tall, much branched, strongly lignified,
± round-topped to irregularly hemispherical, with branches erect to
divergent-ascending or prostrate (typical dwarf espalier shrub -
Hagemann, 1989: 288); wholly glabrous, red or black glands some-
times present in flowers. Stems 3-angled (6-lined) at first, eventually
± terete; cortex green; bark smooth, cinnamon-brown. Leaves all 3-
whorled or uppermost paired, free, petiolate, with petiole 0.4-1 mm
long; lamina (2-)3-8(-10) x 1.2-2.4 mm, oblanceolate to elliptic,
concolorous, glaucous, subcoriaceous: apex rounded, margin
recurved-subindurate, base narrowly cuneate; venation: 1 pair of
laterals sometimes discernible; laminar glands dense to sparse,
scarcely prominent; intramarginal glands dense. Inflorescence 1(-
3)-flowered, terminal and in uppermost leaf axils; bracteoles 1-2.5
mm long, linear, submembranous; pedicels 3-1 1 mm long, slender.
Flowers c. 15 mm in diam.; buds ovoid-pyramidal, subacute. Sepals
green or pink-tinged, (2-)3-4(-5) x 0.75-2 mm, free, not imbricate,
equal, lanceolate to linear, acute chartaceous, margin entire to

145

subentire or minutely denticulate; veins 3(5), not prominent or
branched; laminar glands pale, punctiform, sparse; marginal or
intramarginal glands 1-c. 16, black or reddish, or none. Petals
yellow, sometimes veined red, 7-10(-12?) x 2-3 mm, c. 3 x sepals,
elliptic to oblong, rounded, eglandular or with indistinct pale dots
toward apex. Stamens c. 10, longest 6-8 mm, c. 0.8 x petals; anther
gland (always?) black. Ovary 2-2.5 x 1-1.5 mm, narrowly ovoid-
ellipsoid, acute; styles c. 3 mm long, 1.5 x ovary, curved-ascending.
Capsule 5-6 x 3-3.5 mm, ovoid-conic, narrowly acute,
subcoriaceous, with valves longitudinally vitiate, exceeding sepals.
Seeds blackish, c. 1.7 mm long, narrowly carinate; testa linear-
foveolate.

In fissures of limestone or conglomerate rocks and cliffs, usually in
the Cedrus or Pinus woodland belt; sea level and 1600-1900 m.

Turkey (Pisidian Taurus, Pamphylia).

TURKEY. Antalya: distr. Antalya, Insel Granbusa (Sula Ada), 0 m, 30
May \950(f\),Huber-Morath 13825 (BASBG); distr. Finike, Aykirka, 1950,
Heilbronn & Atilla s.n. (ANK n.v.); distr. Gebiz, Bozburun Dag between
Bogaz Azzi and Tozlu Cukur yayla, 1600m, 24 July 1949 (fl), Davis 15514
(E, K). Isparta: distr. Sut^uler, W. side of Sarp Dag, 1700 m, 29 July 1949,
Davis & BilgerD.\5n\ (E, K).

H. ternatum is a morphological link between the Socotran species
and H. pallens, although the apparently constantly 3-whorled ar-
rangement of the leaves differentiates it from both. It is known as yet
only from (i) two adjacent mountains on the border of vilayets
Antalya and Isparta (Bozburun dagand Sarp dag) and (ii) a small
island off the coast and another, mainland, locality, both near Finike,
south of Tahtali dag. It seems idle to speculate on the significance of
this bitopic distribution, at least until more data are available.

5. Hypericum pallens Banks & Solander in Russell, Aleppo 2nd
ed. 2: 270 ( 1 784); Eig in / Bot. Lond. 75: 1 88 ( 1 937); Poulter in

Map 22 Sect. 23: 5. H. pallens (part) all records

146

N.K.B. ROBSON

Notes R. hot. Gdn Edinb. 21: 181 (1954); N. Robson in P. Davis,
Fl Turkey 2: 369, f. 11/7 (1967), 10: 361 (1988); Mouterde,
Nouv. Fl. Liban Syrie 2: 521, t. 225 f. 2 (1986); I. Hagemann in
Flora 183: 284, if. 58, 59, 62, 63 (1989); N. Robson in Cullen et
al., Eur. Gdn FL 4: 70 (1995). Type: Turkey/Syria, monies inter
Aleppo et Antiocham, 1771 (fl), P. Russell (BM!-holotype).
Fig. 2 ID, Maps 22, 23.

H. cuneatum Poir., Encycl., Suppl. 3: 699 (1814); Choisy, Prodr.
Monogr. Hyperic. : 50 ( 1 82 1 ), in DC., Prodr. 1 : 549 ( 1 824); Spach
inAnnls Sci. not. (Bot.) II, 5: 357 ( 1 836); Jaubert & Spach, ///. pi.
orient. 1: 49, t. 25 (1842); Boiss., Fl. orient. 1: 794 (1867),
Suppl.: 127 (1888); Post, Fl. Syria: 171 (1896), op. cit., 2nd ed.
1: 230 (1932); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd
ed 21: 178 (1925); Boul., Fl. Liban: 68, t. 74 f. 4 (1930);
Stefanoff in God. Agr.-les. Fak. Univ. Sofiva 10: t. 3 f. 17 (1932),
11: 152 (1933), 12: 83 (1934), in Pflanzenareale IV, 1: Karte 2b
( 1 933); Ashberry, Min. trees & shrubs: 1 33 & photograph ( 1 958);
Rech. f. \r\Ark. Bot. 5: 291 (1960); Ingwersen, Man. Alpine pis:
209 ( 1 978). Type: Syria, prope Nourieh, de Labillardiere s.n. (FI-
holotype).

H. myrtilloides Fenzl, Pug. pi nov. Syr.: 1 (1842). Type: Turkey
[Hatay], prope Svedie [Souedieh], Kotschy 108 (W-holotype; K!).

H. tenellum Kotschy ex Boiss., Fl. orient. 1: 794 (1867), in synon.

H. cuneatum [var.] b. fragile Post, Fl. Syria: 171 (1896). Type:
Lebanon, Jabal Fughri, Post s.n. (BEI-holotype).

H. cuneatum [var.] c. pallidum Post, Fl. Syria: 171 (1896). Type:
Lebanon, Nahr-ul-Kalb, Post s.n. (BEI-holotype).

H. cuneatum var. maximum Post, Fl. Syria 2nd ed. 1: 232 (1932),
nomen.

Icon: Jaub. & Spach, ///. pi. orient. 1: 49, t. 25 (1842).

Shrublet 0.05-0.25(0.4) m long, much branched, wiry to lignified,
with branches slender, ascending to prostrate, diffuse; wholly gla-
brous; red or black glands present in leaves, flowers and sometimes
stems. Stems 4-lined and ancipitous at first, eventually terete; cortex
becoming bright red, sometimes with a few black glands; bark
striate, reddish brown. Leaves paired, free, petiolate, with petiole c.
0.5-1 mm long; lamina (2.5-)4-14(-22) x (l-)2.5-7.5(-9) mm,
oblong or elliptic (or rarely ovate) to obovate or oblanceolate,
concolorous, glaucous beneath or on both sides, (sub?)coriaceous;
apex rounded, margin plane to subrecurved, base cuneate to long-
angustate or pseudopetiolate; venation: 0-2(-6) pairs of laterals,
with tertiary reticulation visible in larger leaves; laminar glands
sparse or absent, occasionally black; submarginal glands scattered,
black; intramarginal glands dense. Inflorescence l(2-3)-flowered,
terminal and often also 1 -flowered axillary or terminating branches
from up to c. 6 axils below, sometimes with up to 4 flowering
branches from scattered lower nodes, the whole appearing
racemiform; bracteoles narrowly oblong or absent; pedicels 5-9 mm
long, slender, 'erect or resupinate'. Flowers 10-13(-15) mm in
diam.; buds cylindric-ellipsoid, rounded, bright sealing-wax red (at
least in prostrate form). Sepals green or pinkish red, (2-)3-5(-6) x
0.7-2.5 mm, free or almost so, not or slightly imbricate, equal,
lanceolate or narrowly oblong or ± narrowly elliptic to linear, acute
to rounded, chartaceous to submembranous, margin entire or irregu-

larly glandular-denticulate; veins (3)5-7, not or only midrib promi-
nent, often ± branched; laminar glands pale, punctiform or elongate,
dense to sparse, often also 1-5 black, scattered; inframarginal glands
pale, dense to sparse; marginal black glandular teeth 0-c. 10,
elongate. Petals bright yellow, tinged bright red outside in bud, 6-12
x 2.5^.5 mm, 2-2.5 x sepals, elliptic to oblong, obtuse with
apiculus acute, lateral, with laminar glands scattered, punctiform or
rarely striiform or linear, pale or sometimes some towards apex
black or reddish. Stamens 20-30, longest 6-8 mm, 0.75-0.8 x
petals; anther gland black. Ovary 2 x 1.5 mm, ovoid-ellipsoid to
ellipsoid, acute to acuminate; styles 3-4 mm long, c. 2.5 x ovary,
divergent. Capsule 4-5 x 3^4 mm, ovoid, subcoriaceous, surrounded
by old petals, with valves longitudinally vittate, shorter than sepals.
Seeds reddish brown, c. 1 mm long, ecarinate; testa linear-foveolate.
2n= 16(Reynaud, 1973).

In fissures of hard limestone rocks; 50-1700 m.

Turkey (eastern Cilicia, Amanus), Syria (Latakia), Lebanon (north-
west coastal plain and Lebanon range).

TURKEY. Konya: distr. Ermenek, Hamitseydi bo^ar, between Sarawadi
and Beskuyu rocks, 1500-1700 m, 16 August 1949 (fl), Davis 16232 (E, K).
Icel: distr. Mut?, c. 9 kmW. Abzweigung Silifke-Gulnar, 150m, 4 July 1977
(st), Sorger 77-21 -7 (W); distr. Anamur,Anamur-Gilindire, 50m, 1956 (fl),
Davis & Polunin D.25968 (BM, E, K); Cilicia, Gysel Dere, 300 m, May 1 896
(fl), Siehe 200 (BM, E, JE, K, S); distr. Mersin, Mersin - Kuzucubelen, 1 km
from Kuzukubelen, 600 m, 1 8 June 1 950 (fl), Huber-Morath 9539 (BASBG).
Seyhan: 20 km NW Kadirli, 250 m, 31 May 1973 (fl), Sorger 73-10-11
(BM). Hatay: 4 hrs S. of Antioch [Antakya], 11 June 1884, Post s.n. (BM,
BEI*); Amanus, infra pag. Bityas, c. 200 m, 25 May 1933 (fl), Samuelsson
5418 (S).

SYRIA. Recorded from Latakia (coast), the base of Mt Cassius [Jebel
Akra] (Turkish border) and W. of Sarmada, Sourate (inland), fide Mouterde
(1970:522).

LEBANON. NahrelKelb, 15 May 1942(fl),Davw6102(BM,K);Djier
el Madfour (inter Batroun [Batrun] et Tripoli), 8 June 1932 (fl), Samuelsson
2209 (S); in rupibus regionis ad Brummana, 600-700 m, 7 July 1897 (fl),
Bornmuller 238 (BM, E, G, JE, K).

H. pattens is less woody and more procumbent to prostrate in habit
than H. ternatum, and its area of distribution lies wholly to the east
of that of the latter. The petals are red-tinged and the leaves (always
paired) more variable - usually similar but sometimes broader or
much larger ('var. maximum', from '4 hours south of Antioch', 1 1
June 1884, Post), but none of this variation warrants subdivision of
the species. The inflorescence, although often racemiform, consists
of 1 (rarely 2-3)-flowered sylleptic branches and thus, as Hagemann
(1989: 285) has pointed out, is essentially similar to the 1 -flowered
synflorescence of e.g. H. revolutum (sect. 1 . Campylosporus).

Sect. 24. HETEROPHYLLA N. Robson in Notes R.
hot. Gdn Edinb. 27: 185 (1967) ['Heterophyllum'].

Shrublet, erect, up to 0.25 m tall, semi-deciduous, glabrous, without
dark glands; branching effectively pseudo-dichotomous. Stems 2-
lined to terete, glandular; cortex green; bark smooth, reddish brown.
Leaves opposite, decussate, sessile, free, deciduous at basal articula-
tion, differing in form in lower (perennating) and upper (deciduous)
parts of stem; lamina entire, with venation pinnate and open or 1 -
nerved, without tertiary reticulation; laminar glands linear to
punctiform; marginal gland dots dense; ventral glands absent. Inflo-
rescence 3-12-flowered, with branches dichasial, from 1-3 nodes,
sometimes with lower flowering branches; bracts and bracteoles ±
reduced. Flowers stellate, homostylous. Sepals 5, free, persistent,
erect in fruit, with margin entire; laminar glands linear to striiform;
marginal, submarginal and inframarginal glands absent. Petals 5,

STUDIES IN HYPERICUM

147

persistent, erect but not twisting after flowering, with apiculus
subterminal or obsolete; margin entire; marginal glands absent;
laminar glands linear. Stamen fascicles 3 (i.e. united 2+2+1), dis-
tinct, persistent, with stamens 35-45; filaments basally united;
anthers yellow, gland amber; pollen type X. Ovary with 3 subaxile
placentae, each 2-ovulate; styles 3, free, bases contiguous; stigmas
small. Capsule 3-valved, chartaceous, with valves longitudinally
vittate. Seeds not seen mature, carinate; testa foveolate.

BASIC CHROMOSOME NUMBER (X). 9; ploidy 2.

HABITAT. Clearings inPinus nigra woodland, on limestone; 1 200-
2000m.

DISTRIBUTION. Turkey (north-west and west-central Anatolia).
1 species.

1 . Hypericum heterophyllum Vent., Descr. pi. nouv. : t. 68 ( 1 800);
Choisy, Prodr. Monogr. Hyperic.: 49 (1821), in DC., Prodr. 1:
549 ( 1 824); Spach in Annls Sci. nat. (Bot.) 5: 357 ( 1 836); Boiss.,
Fl. orient. 1: 793 (1867); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 178 (1925); Stefanoff in Kew Bull.
1931: 30 (1931), in God. Agr.-les. Fak. Univ. Sofiya 10: t. 2 f. 8
(1932), 11: 145 (1933), 12: 82 (1934), in Pflanzenareale IV, 1:
Karte 2a ( 1 933); N. Robson in P. Davis, Fl. Turkey 2: 369 ( 1 967),
10: 361 (1988); I. Hagemann in Flora 183: 252", 294, ff. 23-28
(1989). Type: cult, in nursery of J.M. Cels, near Paris, c. 1800
(fl), ex 'Persia'; 'trouve en Perse par Bruguiere et Olivier et
introduit chez Cels en 1'an 6' (i.e. 1797-1798). Neither Boissier
(1867) nor Stefanoff (1931-1934) seems to have doubted
Ventenat's attribution of this species to the Iranian flora, despite
the absence of subsequent collections from that country. Stefanoff
(1931) drew attention to the collection by Whittall (K) labelled
'Smyrna', but correctly treated this locality asWhittall's address.
Later (\933a: 145, 1933ft), he accepted the Smyrna label for
want of a more accurate locality. More recent collections have
revealed the real native region of H. heterophyllum to be north-
western and west-central Turkey, from Balikesir and Afyon to
Ankara and Cankiri, a region traversed by Olivier and Bruguiere
on their journey to and from Iran in 1797-98.

Fig. 22B, Map 23.

Icones: I. Hagemann in Flora 183: 253, f. 24 (1989); Vent., Descr.
pi. nouv.: L 68(1800).

Shrublet (0.05)0.1-0.2(0.3) m tall, much branched to form flattish-
topped bush, with basal lignified parts branched effectively
pseudo-dichotomously, with branches erect and straight to decum-
bent and ± twisted; wholly glabrous, without dark glands. Stems
2-lined at first with short internodes and bearing small scale-like
leaves, later 2-lined to terete with elongate internodes, the latter
(inflorescence-bearing) part obscurely glandular and withering and
leaving the over-wintering basal part bearing pairs of strobiliform

condensed shoots; cortex green; bark smooth, reddish brown to
greyish brown. Leaves free, sessile; perennating scale-leaves 0.5-
1.5 mm long, broadly ovate or orbicular to obovate, cucullate,
obtuse, apiculate to muticous, (all?) elongating in spring to spathulate,
glaucous above, soon deciduous; foliage leaves 5-15 x 1-2 mm,
narrowly elliptic-oblong to linear, concolorous, thinly glaucous
above at least when young, coriaceous, apex acute, margin plane,
base cuneate. Inflorescence (l-)3-5-flowered, terminal and often
with single flowers or 3-5-flowered cymules in axils of 1-2 lower
nodes, the whole up to 13-flowered, rounded-pyramidal to
subcorymbiform; bracteoles triangular-lanceolate to linear; pedicels
very short or absent. Flowers 8-12 mm in diam.; buds ellipsoid,
subacute. Sepals green, 2-3.5 x 0.8-1.2 mm, imbricate, unequal,
oblong to ovate-lanceolate, acute to subacute, entire; veins 5(3), not
or only midrib prominent, unbranched; laminar glands linear to
striiform. Petals bright yellow, not tinged red, 5-8 x 2.5-3 mm, 2.2-
2.5 x sepals, obovate to oblanceolate, obtuse to rounded with
apiculus small, subterminal or obsolete, with laminar glands linear.
Stamens 35^45, longest 4.5-7 mm,c. 0.9 x petals. Ovary c. 1 .5 x 0.7
mm, cylindric-ellipsoid to narrowly ovoid-cylindric, truncate; styles
c. 4 mm long, c. 2.4 x ovary, widely spreading. Capsule 6-8 x 3-3.5
mm, ovoid-cylindric to narrowly cylindric, chartaceous, surrounded
by old petals, longer than sepals, with valves longitudinally vittate.
Seeds 2 on each placenta, not seen mature, rather prominently
carinate; testa foveolate. 2n = 18 (Reynaud, 1973).

Dry clearings in Pinus nigra subsp. pallasiana (Lamb.) Holmboe
woodland and in garigue with Cistus laurifolius L.; 1200-2000 m.

Turkey (north-west and west-central Anatolia).

TURKEY. Balikesir: Dorsunbey, Alagam Intifa, 1200 m, 17 July 1951
(fl), Guresin s.n. (ISTO). Bolii: Passhohe sudlich von Gerede, 1600 m, 8
August 1963 (fi),Huber s.n. (BM,WB). Cankiri: NW of Kizilchamam, 1520
m, 5 September 1966 (fl & fr), Archibald 3316 (E, K). Ankara: Ankara -
Istanbul road opposite turning to ^am^oru, near Kizilcahamam, 19 August
1970 (fl), Fraser- Jenkins 2091 (BM); Ankara - Bolii, 34 km sudostlich
Gerede, c. 1600m, 8 June 1962(e. fl),S0rg*rT62/62/20(E). Afyon: a 20 km
a Test d' Afyon6, 1970 (fl), Contandriopoulos & Quezel 70-534 (MARS).

H. heterophyllum is systematically isolated in Hypericum. It seems
to share with H. aegypticum (sect. 25) an origin near//, balfourii or
H. socotranum (sect. 1) but to be neither wholly apomorphic nor
wholly plesiomorphic in relation to it. It lacks the specializations
that, in Hypericum, are unique to sect. 25 Adenotrias, viz. the
heterostyly syndrome and the carunculate seeds; but it is more
specialized than H. aegypticum in habit, in its biovulate placentae
and in its chromosome number (2n = 1 8, whereas H. aegypticum and
H. russeggeri, both sect. 25, have 2n = 20).

A detailed account of the growth form and behaviour of H.
heterophyllum, both in nature and in cultivation, has been given by
Hagemann (1989). For its relationship with H. aegypticum see p. 84.

Sect. 25. ADENOTRIAS (Jaub. & Spach) R. Keller in
Engl. & Prantl, Nat. Pflanzenfam. 3(6): 209 (1893).

Shrubs and shrublets, erect to prostrate, up to 2 m tall, glabrous,
without dark glands; branching lateral. Stems 4- to 2-lined, glandu-
lar; cortex green, bark smooth, grey-brown to pale grey. Leaves
opposite, decussate, sessile, free, deciduous at basal articulation,
homomorphic; lamina entire, with venation pinnate and open or 1-
nerved, without tertiary reticulation; laminar glands punctiform;

6 Mme Reynaud has kindly informed me that the locality in Kenya cited by her
(Reynaud, 1973: 2 10) is erroneous. She did, however, allude to the correct locality in the
same paper (p. 203).

148

N.K.B. ROBSON

Map 23 Sect. 23: 4. //. ternatum •; 5. //. pattens (part) •. Sect. 24: 1. //. heterophyllum Q Sect. 25: 3. //. russeggeri A, extinct? A (see also Map 24).

marginal glands dense to rather sparse; ventral glands absent. Inflo-
rescence 1-8-flowered, with branches dichasial, from terminal node
only or also from lower nodes, sometimes with a 'sterile' intermedi-
ate zone; bracts and bracteoles reduced. Flowers hypocrateriform to
almost tubular, heterostylous.Sepa/s 5, free, persistent, erect in fruit,
with margin entire; veins 5-9; laminar glands linear; submarginal
glands present; marginal and inframarginal glands absent. Petals 5,
persistent, erect and apically twisting or deciduous after flowering,
the apiculus absent, with basal entire ligulate appendage; margin
entire; marginal and laminar glands absent. Stamen fascicles 3 (i.e.
united 2+2+1), dimorphic, distinct, with stamens 9-c. 48 (3-20 per
fascicle); filaments united to above middle; anthers yellow, gland
amber; pollen type V. Fasciclodes 3, subglobose, alternating with
fascicles. Ovary with 3 axile placentae, each °°- or 2-ovulate; styles
3, dimorphic, bases contiguous; stigmas rather narrowly capitate.
Capsule 3-valved, subcoriaceous, with valves finely longitudinally
vittate. Seeds cylindric, ecarinate, with distal caruncle; testa finely
linear-rugulose or linear-foveolate to linear-scalariform.

BASIC CHROMOSOME NUMBER (x). 10; ploidy 2.
HABITAT. Limestone rocks, often coastal; 0-1000 m.

DISTRIBUTION. South Morocco, Algeria (southern Atlas Mts),
Lampedusa, Malta, Sardinia, Libya (Cyrenaica), Greece (Ionian
Islands, Peloponnisos), Crete, Turkey (Amanus), Syria.

3 species (+ 2 subspecies).
Key to sect. 25. Adenotrias

1 Inflorescence 1 -flowered, sessile; petals and stamens persistent;
ovules numerous in each loculus; leaves elliptic to oblong (1.
aegypticum) 2

Inflorescence ( 1 )2-9-flowered, pedunculate; petals and usually sta-

mens deciduous; ovules 2 in each loculus; leaves narrowly
oblanceolate to linear 4

2(1) Leaves narrowly elliptic or narrowly oblong-elliptic, (7-)
9-18 mm long, sessile; plant erect, (0. 15-)0.3-2 m tall
la. aegypticum subsp. maroccanum

Leaves broadly elliptic or narrowly oblong, 3- 1 0 mm long, subsessile
to shortly petiolate; plant erect to spreading, 0.05-0.6 m tall 3

3(2) Leaves subsessile or to c. 0.3 mm petiolate, plane or subcucullate,
4-10 mm long; plant erect or loosely spreading, 0.04-0.6 m tall
Ib. aegypticum subsp. webbii

Leaves 1-2 mm petiolate, ± incurved-cucullate, 3-6 mm long; plant
spreading, 0.15-0.18 m tall .. Ic. aegypticum subsp. aegypticum

4( 1 ) Peduncle 1 0-20 mm long; leaves 1 .5-3 mm wide; inflorescence 3-
9-flowered; plant 0.1-0.3 m tall, erect to prostrate .. 2. russeggeri

Peduncle c. 2 mm long; leaves 0.6-1.4 mm wide; inflorescence
(l)2-3-flowered; plant c. 0.05-0.06 m tall, prostrate

... 3. aciferum

I . Hypericum aegypticum L., Sp. pi. : 784 ( 1 753), op. cit., 2nd ed.:
1 103 (1763), Hypericum: 6, f. 3 (1776), Amoen. acad. 8: 323, t.
8 f . 3 (1785) ['aegyptiacum']; Murray, Syst. veg. 13th ed.: 583
(1774) ['aegyptium'];Lam.,Encycl. 4: 162(1797);Willd.,5p.p/.
3: 1467 (1802); Ker in Bot. Reg. 3: t. 196 (1817); Choisy, Prodr.
monogr. Hyperic.: 49 (1821), in DC., Prodr. 1: 549 (1824);
Spreng., Syst. veg. 3: 334 (1826) ['aegyptiacum']; Margot &
Reuter, Essai Fl. lie Zante: 35 ( 1 838) [' aegyptiacum']; Chaub. &
Bory, Nouv.fl. Pelop.: 53 (1838); Trevir., Hyper, sp. animadv.: 9
(1861); Nyman, Syll. fl. Eur. Suppl.: 222 (1865); Hook. f. in
Curtis 's hot. Mag. 106: t. 648 1 ( 1 880); Fiori & Paol., Fl. Italia 1:
385 (1898), Iconogr.fl. ital. III.: 143, f. 1247 (1899); Fiori, Nuov.
Fl. Italia 1: 519 (1924) ['aegyptiacum']; Borg, Descr.fl. Maltese

STUDIES IN HYPER1CUM

isi: 247 (1927); Braun-Blanquet & Maire in Bull. Sot: Hist. nat.
Afr. N. 22: 107 (1931) ['aegyptiacum'}; Jah. & Maire, Cat. PI.
Maroc 2: 482 (1932) ['aegyptiacum']; Quezel & Santa, Nouv. Fl.
Algerie 2: 482 (1963); Arrig. in Webbia 20: 324 (1965)
['aegyptiacum']; N. Robson in Tutin et al., Fl. Europaea 2: 264
(1968); Bean, Trees & shrubs hardy in Br. Isles 8th ed. 2: 407
(1973); Ornduff in Bat. J. Linn. Sac: 71: 51 (1975), in Heredity
42: 271 (1979); Ali in Ali & Jafri, Fl. Libya 2: 4, f. 3 (1976);
Haslam, Sell &Wolseley,R Maltese Is.: 199(1977); I. Hagemann
in Bot. Jb. 102: 247 (1981); Pignatti, Fl. Italia 1: 345 (1982);
Greuter in Willdenowia 14: 279 (1984); Reynaud in Adansonia
7B: 90, tt. 1,2 ff. 5, 6 (1985); Greuter, Burdet & Long, Med-
Checklist 3: 263 ( 1 986); Turland in Q. alp. Gdn Soc. 58: 3 1 1 , 3 1 8
& photograph ( 1990); Turland, Chilton & Press, FL Cretan area:
93, map 673 (1993); N. Robson in Cullen et al., Eur. Gdn Fl. 4:
70, ff. 11.2, 11.7, 11.12 (1995). Type: [Libya?] 'Habitat in
Aegypto, D. B. Jussiaeus" in Herb. Juss. 1 1 803 (P-JUS!-lectotype,
selected here). This species occurs in Libya (Cyrenaica) but not
Egypt. The specimen in Paris seems to be the only possible type
material known - there is nothing relevant among Linnaeus's
correspondence with Bernard de Jussieu at the Linnean Society.
An examination of the microfiche shows that this specimen
would appear to be from the Libyan population; and, as it is said
to come from Egypt, I have designated it as the lectotype.
Fig. 22A (subsp. maroccanum), Map 24.

H. creticum Hort. ex Link, Enum. hort. berol. alt. 2: 276 (1822), in

synon.
Elodeaaegyptica (L.) Jack, Mai. Misc. 2(7): 25 (1822) ['Egyptiaca],

in Calcutta J. nat. Hist. 4: 21 1 (1844).
Triadenia microphylla Spach mAnnls Sci. nat. (Bot.) II, 5: 173, t. 4

(1836), Hist nat. veg. Phan. 5: 371 (1836), op. cit. Atlas: t. 1 19 f.

1 (1846). Type as for Hypericum aegypticum.
Episiphis parvifolia Raf., Fl Tellur. 3: 78 (1837). Type as for

Hypericum aegypticum.
Triadenia aegyptica (L.) Boiss., Fl. orient. 1: 783 (1867), Suppl.:

125 (1888) ['aegyptiaca']; Batt. &Trabut, Fl. Algerie Tunisie: 76

(1904); Pamp., Prodr. fl. ciren.: 320 (1930).
Hypericum heterostylum Parl., Fl. Ital. 5: 550 (1875); R. Keller in

Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 175, f. 73A-E

149

(1925); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: t. 2 f. 3
(1932), 11: 140 (1933), 12: 81 (1934), in Pflanzenareale 4(1):
Karte la (1933). Type as for H. aegypticum L.

Elodes aegyptica (L.) [Payer, Traite organogen.: 8, t. 1 (1857)
['aegyptiaca'] sine basionymex]Y. Kimura in/ Jap. Bot. 11: 831
(1935), in Nakai & Honda, Novafl.jap. 10: 18 (1951).

Triadenia aegyptica (L.) Boiss. ['aegyptiaca'] var. microphylla
(Spach) Maire (in sched. 1961).

The above synonymy includes all names that are based on the
Linnaean type, even though some of the references are concerned
wholly or partly with subspp. maroccanum or webbii, not subsp.
aegypticum.

Shrub or shrublet 0.05-2 m, tall, much branched, bushy or ±
spreading, with branches erect to divergent or rarely decumbent to
ascending; wholly glabrous, without dark glands. Stems 2 (rarely 4)-
lined and ancipitous at first, soon terete, glaucous, with internodes
shorter than leaves. Leaves free, sessile to shortly (c. 0.5 mm)
petiolate, persistent fore. 2 seasons; lamina (3-)3.5-18 x (l-)1.5-5
mm, narrowly to broadly elliptic or narrowly oblong, concolorous, ±
densely glaucous, coriaceous, apex acute to obtuse, margin plane to
incurved-cucullate, base cuneate; venation: 1 pair of laterals or 1-
nerved, with midrib sometimes pinnately branched and ± prominent
beneath; laminar glands dense; marginal glands dense. Inflores-
cence 1 -flowered, terminal and from up to c. 4 axils below, with
short flowering branches (usually after a 'sterile' region) from up to
c. 1 1 nodes, or 'sterile' region absent and flowers or flowering
branches from up to t: 22 nodes, the whole spiciform, with elongate
branches originating below flowering region; bracts absent or foliar,
clasping calyx, persistent, with glands punctiform and striiform;

Map 24 Sect. 25: 1 . H. aegypticum: a. subsp. maroccanum •; b. subsp. webbii •, also NE Crete; c. subsp. aegypticum A; 2. H. russeggeri A (see also
Map 23); 3. H. aciferum D.

150

pedicels very short or absent. Flowers c. 5-10 mm in diam.; buds
narrowly ovoid-ellipsoid, subacute. Sepals green, (2.5)-3.5-5.5 x
1-2 mm, imbricate, subequal, oblong to elliptic or lanceolate-
elliptic, rounded to apiculate or obtuse, ± cucullate, stiffly erect;
veins c. 9, only midrib prominent, unbranched; laminar glands
linear; submarginal glands rather dense. Petals bright to rather pale
yellow, persistent in fruit, 6.5-12(-14) x 2-3 mm, c. 2 x sepals,
oblanceolate, distally outcurved to form hypocrateriform pseudo-
tubular corolla, rounded, with basal ligulate appendage
oblanceolate-spathulate to narrowly oblong with margin incurved.
Stamens 18-c. 48 (i.e. single fascicle 5-10, double fascicles 5-29),
with filaments in each fascicle c. 0.6-0.7 united, the longest 2.5-6.5
mm (long-styled) or 6-9 mm (short-styled), 0.3-0.5 x petals (long-
styled) or 0.75-0.95 x petals (short-styled), persistent in fruit.
Fasciclodes (lodicules) c. 0.6-0.7 mm long, flat-topped. Ovary 1 .5-
4 x 0.5-1 mm, narrowly ellipsoid to very narrowly ovoid-conic,
acute to truncate; styles 0.5-1 mm (short-styled) or 2.3-3.2 mm
(long-styled), 0.25-0.33 x ovary (short-styled) or 2.25-4 x ovary
(long-styled), erect; ovules °° on each placenta. Capsule 5-7 x 3-3.5
mm, cylindric-ellipsoid to rather broadly ellipsoid, longer than
sepals, with valves longitudinally vittate. Seeds dark brown, c. 1.5
mm long, ecarinate; testa finely linear-foveolate to linear-rugulose;
caruncle often lobed (cf. Reynaud, 1985: 91, t. 2, ff. 5, 6). 2n = 20
(subsp. maroccanum) (Ornduff in Robson, \911a: 334).

In fissures of limestone rocks or on limestone scree in valleys, often
coastal; 30-c. 1600m (Morocco), c. 1000-1 200m (Algeria), 0-500
m (Malta, Lampedusa), 700-800 m (Sardinia), 0-300 m (Ionian
Islands, Peloponnisos), coastal (Crete), 1-750 m (Cyrenaica).

South Morocco (coast, Grande Atlas, Anti-Atlas), Algeria (Atlas),
Lampedusa, Maltese Islands (Malta, Gozo, Comino), Sardinia (south-
east), Ionian Islands (Kefallinia, Zakinthos), Greece (eastern
Peloponnisos), Crete (north-west, north-east), Libya (Cyrenaica
near Derna). Not known from Egypt.

H. aegypticum has a very disjunct distribution from south Morocco
via the southern slope of the Atlas Mts in Algeria, Lampedusa,
Malta, Sardinia, the Ionian lalands, the Peloponnese and Crete to
Cyrenaica, the whole forming a morphological and geographical
cline in the direction indicated. Thus the Moroccan and Algerian
plants are erect (to 2 m) with long sessile leaves and large flowers,
the island and Peloponnese plants are erect to spreading with
intermediate length, sessile to subpetiolate leaves and intermediate-
sized flowers, and the Libyan plants are low and spreading with
small petiolate leaves and small flowers. Throughout this wide range
the variation is almost continuous; but it is possible, I think, to
separate an island and Peloponnese form from the African forms at
subspecies level, viz. as subspp. maroccanum, webbii and
aegypticum.

H. aegypticum is the most primitive species in sect.Adenotrias, its
nearest relative apparently being a Socotran member of sect. 1.
Campylosporus, H. balfourii. It differs from the Socotran plant in
size of leaf and flower, but also particularly in the heterostyly and
other adaptations for specialized insect pollination (pseudo-tubular
flowers, petal appendages, fasciclodes acting like grass lodicules by
swelling to open the flower, stamen filaments united) and dispersal
(carunculate seeds). The narrow leaves of the primitive (Moroccan)
form are more similar to the those of//, balfourii than to those of the
latter's close relative, another Socotran endemic, H. socotranum.

la. Hypericum aegypticum subsp. maroccanum (Pau) N. Robson
inBull. nat. Hist. Mus. Lond. (Bot.)23: 68 (1993). Type as for//.
aegypticum var. maroccanum Pau.

N.K.B. ROBSON

H. aegypticum sensu Braun-Blanquet & Maire in Bull. Soc. Hist,
nat. Afr. N. 22: 107 (1931) ['aegyptiacum']; Jah. & Maire, Cat.
PL Maroc 2: 482 (1932); et auct. maroc. et alger. plur.

H. aegypticum var. maroccanum Pau in Cavanillesia 4: 157 (1932)
['maroccana'] (Spanish); Maire in Bull. Soc. Hist. nat. Afr. N. 24:
206 (1933) (Latin). Type: Morocco, Sud ouest, Cap Ghir, versant
Sud, 27 March 1931 (fl & fr), Jahandiez 57 (BC-holotype; BM!).

Plant erect, (0.15-)0.3-2 m tall, with branches erect to ascending.
Leaves sessile, plane; lamina (7-)9-18 x (2-)3-4 mm, narrowly
elliptic or narrowly oblong-elliptic, acute. Sepals 5-6 mm long.
Petals 10-12(-14?) mm long.

Morocco, Algeria.

MOROCCO. Sud Ouest: between Agadir and Cap Ghir, 30 m, 1 April
1972 (fl), Davis 53959 (BM); 30 km N. of Agadir, 10 km up ImmouzerValley,
300 m, 2 February 1974 (fl), Miller, Russell & Sutton 472 (BM, RNG*).
Grande Atlas: Immouzerdes-Ida-Outanane to Oulma, 200 m, 19 March 1969
(fl & fr), P.M. & J. Davis D.48463 (BM). Anti Atlas: au dessus de Taliouine,
1500-1600 m, Maire 1397 (MPU*).

ALGERIA. Atlas Saharien: Mts des Ksour, Djebel Grouz; Djebel Bou
Kahil, Ksar Kahil. Both records fide Quezel & Santa (1963: 682).

Subsp. maroccanum is confined to the southern slopes of the Atlas
Mountains and their foothills down to the Atlantic Ocean. In view of
the almost strictly maritime distributions of the other subspecies, it
is tempting to suggest that the southern edge of the Atlas is an
ancient coastal region. Perhaps, as Melville (1967: 293) has sug-
gested, the ancient Tethys Sea was to the south of the Atlas, not, as
most workers still believe, to the north of it.

Ib. Hypericum aegypticum subsp. webbii (Spach) N. Robson in
Bull. nat. Hist. Mus. Lond. (Bot.) 23: 68 (1993). Type as for
Triadenia webbii Spach.

H. maritimum Sieber, Reise Kreta 2: 322 (1823); C.B. Presl in
Oken, Isis 21: 275 (1828); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 175 (1925); Stefanoff in God. Agr.-les.
Fak. Univ. Sofiya 11: 140 (1933), 12: 81 (1934).Type: Crete, Cap
Maleca, Perivolizza [Perivolitsa], 1817 (fl), Sieber s.n. (W?-
holotype; H!).

Triadenia webbii Spach in Annls Sci. nat. (Bot.) II, 5: 174, t. 5A
(1836), Hist. nat. veg. Phan. 5: 372 (1836), Atlas: t. 119 f. 2
(1846); Hal., Consp. Fl. Graeca 1: 285 (1900). Type: Malta, 'In
rupestribus insulae Melitae', Webb s.n. (Fl-holotype).

T. thymifolia Spach in Annls Sci. nat. (Bot.) II, 5: 174, t. 5B (1836),
Hist, nat veg. Phan. 5: 372 (1836), Atlas: t. 119 (1846). Type:
Malta, 'In horto insulae Melitae' [Jardin d'acclimitation de 1' ile
de Make], Webb s.n. (Fl-holotype).

T. sieberi Spach in Annls Sci. nat. (Bot.) II, 5: 175, t. 5C (1836), Hist.
nat. veg. Phan. 5: 373 (1836). Type: Crete: 'In Cretae maritimis',
Sieber (Fl-holotype).

Hypericum spachianum Steud., Nomencl. hot. 2nd ed. 1: 789 ( 1 840).
Type as for Triadenia thymifolia Spach.

H. webbii (Spach) Steud., Nomencl. hot. 2nd ed. 1: 790 (1840);
Nyman, Syll.fl. Eur. Suppl.: 222 (1865); Stefanoff in God. Agr.-
les. Fak. Univ. Sofiya 10: t. 3 f. 3 (1932).

H. sieberi (Spach) Nyman, Syll.fl. Eur. Suppl.: 222 (1865).

Triadenia maritima (Sieber) Boiss., Fl. orient. 1: 784 ( 1 867); Halacsy,
Consp. fl. graec. 1: 284 (1900); Hayek, Prodr.fl. pen. Bale. 1: 530
(1925); Rech. f., Fl. aegaea: 260 (1943).

T. maritima [var.] B. webbii (Spach) Hayek, Prodr.fl. pen. Bale. 1:
530(1925).

T. maritima subsp. weissii Stamatiadou (1971) in sched.

Hypericum aegypticum sensu auct. mult, pro parte excl. typum.

STUDIES IN HYPERICUM

151

Fig. 22 A. H. aegypticum subsp. maroccanum: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) petal ligule; (0 stamen fascicle; (g) half flower showing long
styles, short stamens and 'lodicules'; (h) capsule. B. H. heterophyllum: (i) habit; (j) mature leaf, upper and lower surfaces; (k) condensed axillary shoots;
(1) sepal; (m) petal; (n) capsule (a, i x Vr, g x 2; b, j, k, x 3; c-f, h, 1-n x 4). A. Davis 48463. B. Nyddegger 15521.

152

N.K.B. ROBSON

Icones: Ker in Bot. Reg. 3: t. 196 (1817); Hook. f. in Curtis's hot.
Mag. 106:1.6481 (1880).

Plant erect to loosely spreading, 0.04-0.4(-0.6) m tall, with branches
erect or usually ± spreading and often tortuous, forming bushes up to
1 m across. Leaves subsessile to shortly (c. 0.3 mm) petiolate, plane or
subcucullate; lamina 4-10 x 1.5-3 mm, narrowly oblong to broadly
elliptic, acute to obtuse. Sepals 5-6 mm long. Petals 8-14 mm long.

Lampedusa, Malta, Sardinia, Ionian Islands, Greece (Peloponnisos),
Crete.

LAMPEDUSA. Lampedusa, April 1884 (fl), Lojacono 96 (BM, K),
May 1898 (fr), Ross 1 17 (JE, K).

MALTA. Malta: Oued Babur, S. of Qrendi, 50-100 m, 10 March 1970
(fl), Davis 49443 (K); Dingli, 1926 (fl), Bankart s.n. (BM). Comino: fide
Borg (1927: 247) and Haslam, Sell & Wolseley (1977: 199). Gozo: Wied
Xlendi, 21 March 1872 (fl), Duthie s.n. (BM).

SARDINIA (Central). Nuoro: valley of R. Flumendosa between Seulo
and Villanovatulo, 700-800 m,fide Arrigoni (1965).

IONIOI NISOI. Kefalinnia [Cephalonia]: [ Argostoli] HagiosTheodoros,
10-15 May 1926 (f\),Bornmuller 283 (BM,JE, K).Zakinthos [Zante]: Agios
Georgios nearVolimes, 0-300 m, 1 1 January 1940 (fl), Davis 1 107 (BM, K);
near Kampi, c. 250 m, 1 April 1980 (fl), Young 630 (H, K).

GREECE. Peloponnisos, Messina: Island of Sphakteria [Sfakteria], S.
extremity, April 1862 (fl), J.S. Mill s.n. (K); distr. Trifilia, NW of Gargaliani
above bifurcation to Marathoupolis, 300 m, 30 April 1971 (fl), Stamatiadou
71 (ATH*,BM).

KRITI. Kidhonia: Peninsula Akrotiri, prope Perivolitsa, 20-50 m, 10
October 1966 (st), Greuter 7706 (G*, K); Maleca (Perivolizza), 1820 (fl),
Sieber s.n. (FR, G-DC, H, JE, K). Sitia: small bay NE of the Faneromani
Monastery, coastal cliffs, 31 March 1982 (fl?). Kalheber 82^56 (Herb.
Kalheber.*, Herb. Greuter.*).

CULTIVATED. Specimens seen from France (1813-1825), Germany
(Berlin, 1818-1888)and England ( 1836-1 99 D.According to Hooker( 1880),
H. aegypticum was introduced into cultivation by Andre Thouin in 1787.

Although H. maritimum (NW Crete) has been separated from the
other European populations of//, aegypticum mainly because of its
spreading habit, this does not provide a differentiating character.
Indeed Greuter (in Greuter, Matthas & Risse, 1984) describes the
plants at the second Cretan locality as 'more normal-looking', and
Turland's photograph (1990: 313) bears this out.

Hooker described plants in cultivation at Kew as hardy when
given a little basal winter protection, and as growing erect in
sheltered locations, observations which agree with my experience
when growing it in east Surrey.

Ic. Hypericum aegypticum subsp. aegypticum
For synonymy (including pro parte) see p. 148.

Plant spreading, 0.05-0.18 m tall, with branches ± spreading and
tortuous, forming low bushes. Leaves shortly (0.4-0.5 mm) peti-
olate, always(?) ± incurved-cucullate; lamina 3-6 x 1-2 mm,
narrowly oblong to broadly elliptic, acute. Sepals 3.5-5 mm long.
Petals 6.5-9 mm long.

Libya (Cyrenaica - Jebel el Akhdar).

LIBYA. Cyrenaica: Wadi Derna, second scarp above Apollonia, 4 April
1939 (fl), Simpson 39291 (BM), Sandwith 2342 (K); 12-15 km W. of Derna
on Gubba road, 300 m, 28 July 1970 (fl), Davis 50231 (K); Messa a ovest di
Cirene, U. Tmista, 29 April 1934 (fl), Pampanini & Pichi-Sermolli 4986
(BM, Fl*, K).

Although the type is said to be from 'Aegypto', there is no likely
habitat for H. aegypticum in the Mediterranean part of Egypt. Lin-
naeus's own figure shows a small-leaved plant that matches those in
the Libyan population. It is thus all but certain that the type comes
from Jebel el Akhdar.

2. Hypericum russeggeri (Fenzl) R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 3(6): 209 (1893), 2nded.21: 175 (1925); Boul.,F/.
Liban: 67 (1930); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya
10: tt.2 f. 2, 3 f. 2 (1932), 11: 140 (1933), 12: 81 (1934), in
Pflanzenareale 4(1): Karte la (1933); N. Robson in P. Davis, Fl.
Turkey 2: 370, ff. 1 1.8, 13.3 (1967); Mouterde, Nouv. Fl. Liban
Syrie 2: 52 1 , t. 224 f. 3 ( 1 970); Reynaud in Adansonia 7B: 88, tt.
1 f. 4, 2 ff. 1-2 (1985); Greuter, Burdet & Long, Med-Checklist
3: 272 (1986). Type as for Triadenia russeggeri Fenzl.

Maps 23, 24.

Triadenia russeggeri Fenzl, Pug. pi. nov. Syrie: 1 (1842), in
Russegger, Reisen 1(2): 907 (1843), Atlas 3: t. 13 (1843); Boiss.,
Fl. orient. 1: 784 (1867); Post, FL Syria 2nd ed. 1: 227 (1932);
Rech.f., Fl. aegaea: 260 (1943). Type: Turkey, 'Hab. in Syria
prope Svedie. et ostia Orontis', 1836 (fl), Kotschy 101 (W!-
holotype; BM!, E!, G!, K!, U!-isotypes). The labels on the
Kotschy and Montbret collections of 1 836 have led to a certain
amount of confusion. Most specimens of Kotschy 101 merely
state 'In monte Tauro'. The Edinburgh specimen, however, has
'Syria, Svedia' (i.e. Turkey, vil. Hatay) and Kotschy 1 1 1 (K) has
'circa Swediam prope Antioch'. It would seem likely, then, that
Svedia is the type locality.

Elodea russeggeri (Fenzl) Walp., Repert. hot. syst. 1: 391 (1842).

Adenotrias phrygia Jaub. & Spach, ///. pi orient. 1: 76, t. 39 ( 1 842).
Types: Turkey, Canakkale, 'crescit in Phrygia, prope Adramiti'
[Edremit] [1836], Montbret s.n. (Fl-lectotype, selected here);
Turkey, 'ad monies Cassii radices, in rupestribus secus Orontem',
[May 1834] (fl), Aucher 872 (P-syntype; BM!, G!, K!-isotypes).

A. kotschyi Jaub. & Spach, ///. pi. orient. 1: 77 ( 1 842). Type: Turkey,
'in Tauro', Kotschy 101 (P-holotype). See note under Triadenia
russeggeri.

Hypericum empetrifolium sensu Kotschy ex Jaub. & Spach, ///. pi.
orient. 1: 77 (1842), in synon. Adenotrias kotschyi.

Elodes russeggeri (Fenzl) Greuter in Candollea 20: 216 (1965).

Icon: Jaub. & Spach, ///. pi. orient. 1: 76, t. 39 (1842).

Shrub or shrublet 0.1-0.3 m tall, much branched, spreading to
prostrate, with branches ascending to prostrate, often ± tortuous;
wholly glabrous, without dark glands. Stems 4-lined and ancipitous
at first, soon 2-lined, glaucous, with internodes shorter to rarely
longer than leaves. Leaves free, sessile or subsessile, persistent until
second season; lamina 4-20 x 1 .5-3 mm, narrowly oblanceolate,
concolorous, ± densely glaucous, coriaceous, apex obtuse, margin
plane or proximally incurved, base cuneate; venation: apparently 1-
nerved, with midrib sometimes prominent beneath; laminar glands ±
dense. Inflorescence 3-7(-9)-flowered, terminal and sometimes
also terminating short shoots or solitary in axils of lowermost
(fallen) leaves; bracts foliar; bracteoles linear, squamiform, decidu-
ous; peduncle 10-20 mm long, pedicels 1.5-2.5 mm long. Flowers
c. 10 mm in diam.; buds narrowly ovoid-ellipsoid, subacute. Sepals
green, 2-3 x 0.7-0.9 mm, imbricate, subequal, narrowly oblong to

STUDIES IN HYPER1CUM

narrowly triangular-lanceolate, subrounded to acute, plane to
incurved, stiffly erect; veins 5(-77), only midrib sometimes promi-
nent, unbranched; laminar glands linear; submarginal glands dense.
Petals rather pale yellow, deciduous, 6-9 x c. 3^.5 mm, c. 3 x
sepals, oblanceolate, distally outcurved to form hypocrateriform
pseudo-tubular corolla, rounded, with basal ligulate appendage
oblanceolate? with margin incurved. Stamens c. 30 (i.e. single
fascicles c. 8, double fascicles c. 10-12), with filaments in each
fascicle c. 0.8 united, the longest c. 3-4.5 mm (long-styled) or 4.5-
5 mm (short-styled), deciduous later than petals. Ovary 1 .5 x 0.7
mm, narrowly ellipsoid, acute; styles c. 1 .5 mm (short-styled) or 2
mm (long-styled), about equalling ovary (short-styled) or c. 1.35 x
ovary (long-styled), suberect; ovules 2 on each placenta one ascend-
ing, one pendent. Capsule 2.5-4 x c. 1-1 .5 mm, cylindric-ellipsoid,
longer than sepals, with valves longitudinally vittate. Seeds blackish
brown, c. 2 mm long, ecarinate; testa finely densely linear-
scalariform; caruncle lobed (cf. Reynaud, 1985: 91, t. 2 ff. 1-2). 2n
= 20(n= 10; Reynaud, 1981).

Among calcareous rocks; sea level -100 m.

Turkey (Hatay; probably extinct in Canakkale); Syria (northern
coast).

TURKEY. Canakkale: Edremit [prope Adramiti], 1836 (fl), Montbret
s.n. (FI*). Hatay: Antakya, d. Samandag, near Cevlik, 100 m, 6 May 1965
(fl), Coode & Jones 636 (E), 636A (K); Monte Cassio [Akra Dag], May 1 834
(fl), Aucher-Eloy 872 (BM, G, K).

SYRIA. Nusairy Mts [Jebel el Ansariye], Banias [Baniyas], 13 April
1885 (fl), Post s.n. (BM); Slerife, route de Nasimata vers Ghab, 1973 (fl),
Reynaud 73.204 (MARS).

H. russeggeri differs from H. aegypticum in (for example) leaf
shape, the deciduous petals and stamens and the 2-ovulate placentae.
It 'continues' the discontinuous eastward cline of that species be-
yond Crete, into Asia Minor and Syria, where it occurs in similar
maritime and submaritime habitats. It would appear to be extinct
near Edremit (if Montbret's locality is accurate) and to be restricted
now to the coast and foothills of the southern Amanus mountains
(Gavur daglari) and Akra Dag round the mouth of the Orontes (Asi).
Boissier's 1846 locality 'ad ruinas Seleuciae' refers to Seleucia
Pieria at the mouth of the Orontes, not to Seleucia (Silifke) on the
coast south-east of Mersin (vil. Icel).

3. Hypericum aciferum (Greuter) N. Robson in Repert. nov. sp.
74: 23 ( 1 967), inTutin et al., FL Europaea 2: 264 (1968); Greuter
\nAnn. Mus. Goul. 1: 39 (1973); I. Hagemann inBot. Jb. 102: 247
(1981); Turland in Q. Bull. alp. Gdn Soc. 58: 310 & fig. (1990),
in Bot. J. Linn. Soc. 108: 351 (1992); Turland, Chilton & Press,
FL Cretan area: 93, map 672, t. 4 f. 2 (1993). Type as for Elodes
acifera Greuter.

Map 24.

Elodes acifera Greuter in Candollea 20: 215, f. 13 (1965). Type:
Crete, prov. Sfakia, ad ostium faucium Domata, 20^0 m, 5 June
1962 (fl & fr), Greuter 4669 (Herb. Greuter.-holotype; BM!, G,
LD, W, Z!-isotypes).The holotype is long-styled; Greuter 4669a
(Herb. Greuter.) is short-styled.

Icon: Greuter in Candollea 20: 215, f. 13 (1965).

Shrublet c. 0.05-0.06 m tall, much branched, prostrate to ascending,
elongate, ± tortuous, distally ascending and bearing short erect to
ascending clustered flowering shoots, forming mats over 600 mm in
diam; wholly glabrous, without dark glands. Stems 2-lined at first,
soon terete, densely glaucous, with internodes shorter than leaves.

153

Q 9

ex

O '

N 1

O- -A

P P

Leaves free, sessile, persistent until second season; lamina 5-12 x
0.6-1.4 mm, narrowly linear-spathulate, aciform, concolorous,
densely glaucous, coriaceous, apex acute to subacute, margin plane
or incurved or lamina subtriquetrous, base narrowly cuneate; vena-
tion: 1 -nerved, with midrib subprominent beneath; laminar glands
distally dense or all scattered. Inflorescence ( 1 )2-3-flowered, termi-
nal and terminating short axillary shoots; bracts foliar, bracteoles
linear-acicular, deciduous; peduncle c. 2 mm long, pedicels 1-2 mm
long. Flowers c. 8 mm in diam.; buds narrowly ovoid, acute. Sepals
green, 3-3.5 x 1 mm, imbricate, equal, narrowly oblong or narrowly
oblong-lanceolate to linear, obtuse, plane to subincurved, suberect;
veins 5, only midrib sometimes prominent, unbranched; laminar
glands linear, submarginal glands rather sparse. Petals bright yellow,
deciduous, c. 7.5 x 1.5 mm, c. 2.5 x sepals, oblanceolate, apically
outcurved to form very short-lipped pseudo-tubular corolla, rounded,
with basal ligulate appendage linear, acute. Stamens 9 (i.e. 3 fasci-
cles of 3), with filaments in each fascicle almost completely united,
the longest 3.5-4.5 mm (long-styled) or c. 1.5 mm (short-styled),
persistent (? or tardily decicuous). Fasciclodes (lodicules) 0.3 mm
long, truncate. Ovary 1 .5 x 1 mm, ovoid, acute; styles c. 3 mm (long-
styled) or scarcely 0.5 mm (short-styled), 2 x ovary (long-styled) or
0.33 x ovary (short-styled), suberect; ovules 2 on each placenta, one
ascending, one pendent. Capsule c. 4 x 2 mm, narrowly ovoid,
longer than sepals, with valves longitudinally vittate. Seeds not seen.

Among calcareous rocks; 5^40 m.

Crete (south-west); known from only two localities.

KRITI. Sfakia, lower part of Domata gorge, near Aj. Rumeli, 20-40 m,
5 June 1962 (fl), Greuter 4669 (BM, Z); Selinos-Sfakia, mouth of Tripiti
gorge, W.-facing side, 26 October 1989 (fl), Turland 105 (BM).

This most reduced member of sect. Adenotrias has a very restricted
relict distribution. It is more prostrate than H. russeggeri (its nearest
relative) and has narrower leaves, fewer flowers, petals that curve
out only at the tip, and (probably) persistent stamen fascicles with
fewer stamens per fascicle. Although it is apparently so rare, its
habitat would seem to be similar to those of H. aegypticum and H.
russeggeri (Turland, 1992: 351).

Sect. 26. HUMIFUSOIDEUM R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 3(6): 21 1 (1893).

Hvpericum sect. Pulogensia N. Robson in Blumea 20: 259 (1973

"['1972']).
Hypericum sect. 9. Hypericum sensu N. Robson in Bull. Br. Mus.

nat. Hist. (Bot.) 5: 320 (1977) pro parte, quoad syn. sect.

Pulogensia, i.e. spp. Taiwan, incl.

Shrubs, subshrubs or wiry perennial herbs, erect to prostrate, up to
1 .5 m tall, the shrubs evergreen, glabrous, usually with dark glands;
branching pseudo-dichotomous and/or lateral. Stems 2— 4(6)-lined
and ancipitous when young, sometimes becoming 2-lined or terete,
eglandular or rarely black-dotted; cortex green or usually wine-red;
bark smooth, red-brown to grey-brown. Leaves opposite, decussate,
sessile or very shortly petiolate, free, in shrubs eventually deciduous

154

N.K.B. ROBSON

at base; lamina entire, with venation pinnate, open or closed, the (ovate or oblong or elliptic to orbicular) or, if oblanceolate to linear

tertiary reticulation absent or rudimentary or rarely rather conspicu- (Spp.6(i),7b, 10 p.p.), then shortly petiolate and/or plant a wiry herb
ous; laminar glands pale, linear to punctiform, ± prominent;

submarginal glands absent; intramarginal glands pale and/or dark, 2(1) Leaves spreading or ascending, (7-)8-12 mm long, plane; styles

dense to rarely sparse; ventral glands absent. Inflorescence 1-flow- (2-) 2.5-4 mm long 1. sewense

ered with branches pseudo-dichotomous or rarely up to 10-flowered

,.,.,, . . -ic ~ Leaves imbncate-appressed, 2-9 mm long, plane to incurved; styles

with branches dichasial/monochasial or mixed, from up to 2 nodes, . c 0 , ,

1.5—2.5 mm long 3

rarely with flowering branches from up to 10 lower nodes; bracts and

bracteoles foliar, rarely somewhat reduced. Flowers stellate or 3(2) Leaves ± incurved, narrowly elliptic or lanceolate-elliptic to linear,

rarely ± obconic (Sp. 4), homostylous. Sepals 5, free, persistent, laminar 8lands most'y linear to st"iform; placentation parietal;

erect to somewhat spreading in fruit, with margin entire or rarely black g|ands near'y always Present at 2least ™ ^"S

(Spp. 6(i) p.p., 7a) irregularly glandular-ciliate; laminar glands pale,

linear to punctiform; submarginal glands absent; inframarginal glands Leaves plane, ovate-lanceolate to elliptic or narrowly elliptic-ob-

or marginal glands pale and/or dark (black or rarely reddish) or long, laminar glands mostly striiform to punctiform; placentation

absent. Petals 5, persistent, spreading after flowering, with apiculus axile; black §lands completely absent 3. macgregorii

subterminal or obsolete or absent; margin entire; marginal glands 4(i) Leaf laminar glands all or mostly linear, sometimes with parallel

absent or few or rarely forming a row (Spp. 6(i) p.p., 7), black rows of dots; inflorescence branching wholly pseudo-dichotomous

(sometimes ± prominent) or rarely pale; laminar glands linear to or with weak I -flowered laterals from 1-2 nodes below; erect shrub

punctiform, pale and/or black. Stamen fascicles 3-5, obscure, or 4. bifurcation

stamens not obviously in fascicles, persistent, with stamens 1 3-80; Leaf ,aminar glands al, or mostly shortly striiform and/or puncti.

filaments basally united or apparently free; anthers yellow, gland form; inflorescence branching various, if wholly

amber or black; pollen type X. Ovary with 3-5(6) parietal (intrusive) pseudo-dichotomous, then plant a diffuse herb 5

to axile placentae, each «>-ovulate; styles free, bases contiguous;

. . . 4 „ , _ c,,. . , 5(4) Inflorescence branching dichasial or mixed dichasial/pseudo-di-
stigmas narrowly or scarcely capitate. Capsule 3-5(6)-valved,

,. chotomous; plant an erect or straggling shrub or snrublet

subconaceous to chartaceous, with valves longitudinally vittate, 5 papuanum

rarely also diagonally vittate (Sp. 6) or almost smooth (Sp. 7b) or

thin-walled, bacciform and indehiscent (Sp. 10). Seeds cylindric, Inflorescence branching pseudo-dichotomous or, if dichasial, then

not or scarcely carinate; testa linear-reticulate to linear-foveolate or Plant a ± diffuse Aizomatous shrublet or suffrutescent herb 6

scalariform or scalariform-reticulate. 6(5) Styles longer than or equalling ovary; fruit capsular 7

BASIC CHROMOSOME NUMBERS (X). 1 2, 9, 8; ploidy 2. Styles shorter than ovary or, if equalling or longer than it (Sp. 10),

then fruit baccate 1 1

HABITAT. Wet to dry upland grassland, woodland clearings and

disturbed ground; 1 500-4300 m (New Guinea), c. 2800 m (Luzon), 7(°) Plant without black glands; leaves ovate to elliptic or oblong, with at

1800-3997 m (Taiwan), 1800-3300 m (Java, Sumatra), 300-3600 m least some striiform g'ands 6- pulogense

(Africa, Madagascar). Plant with at least anther gland black; stems diffuse, wiry; leaves

„.,,,. , . ovate to linear, with glands punctiform or somewhat elongate ... 8
DISTRIBUTION. New Guinea (Papua/New Guinea and Irian Jaya),

Philippines (northern Luzon), Taiwan, Indonesia (Sumatra, Java), 8(7) Flowers 12-30 mm in diam.; inflorescence branches (when present)

Madagascar, Africa (Cape Province to Ethiopia, Sudan Republic, axillary; plant suberect to ascending 9

Zaire and Angola; also Cameroon and Fernando Poo). Flowers 7_,0 mm in diam . inflorescence branches (when present)
1 2 species (+ 1 subspecies) pseudo-dichotomous; plant usually diffuse, ascending to decum-
bent (7. H. beccarii) 10

N.B. At a late stage in the work on Part 6, 1 realized that my original

,Jr__. , . _ . . 9(8) Sepals without black laminar streaks or, if with them, then apex

idea (Robson, 1973) that the relationships of two Taiwan species obtuse; ,eaves yariable jn size and shape but .f under 15 mm ,ong

were with H. pulogense Merr. ( from Luzon) was correct, and that then narrow (l:b = 4-12) 6(i) H. nagasawai

they were not, as I had subsequently concluded (Robson, 1977a),

related to a Japanese species of sect. 9. Hypericum, H. yezoense SePals with black laminar streaks< aPex acute to subacuminate;

Maxim. Indeed, the shrublet or wiry herbaceous habit of these two leaves short (4~n mm lon*> and broad .<'* '<-*•»

6(11) H. nokoense

species would be quite anomalous in sect. Hypericum. H. nagasawai

Hayata and H. nokoense Ohwi have therefore had to be inserted in 10(8) Leaves broadly oblong to elliptic-oblong or narrowly

the numerical sequence of sect. Humifusoideum as Spp. 6(i) and 6(ii) obovate; capsule densely and prominently vittate

respectively 7a' beccarii subsp. beccarii

Leaves oblanceolate to linear; capsule sparsely and obscurely vittate
Key tO Sect. 26 Humifusoideum (almost smooth) 7b. beccarii subsp. steenisii

1 1(6) Fruit capsular, usually erect; styles usually 3-4 12

Flowers solitary, with branching almost always wholly lateral;

leaves relatively narrow (lanceolate or narrowly elliptic to linear), Fruit baccate, on reflexed pedicel; styles 4-5 ... 10. peplidifolium

sessile; shrub or shrublet 2

12(11) Stems erect, stout; leaves usually sessile 8. natalense

Flowers in dichasia with branching lateral or, if solitary, then

.... 7 I-, Stems decumbent to ascending, slender; leaves shortly petiolate
branching pseudo-dichotomous' or basal; leaves relatively broad

.... 9. M ins

7 I.e. with (usually) paired axillary branches continuing growth from uppermost node
of flowering branch.

Hypericum sewense N. Robson in Blumea 20: 254 (1973), in
Steenis, Fl. Males. 1, 8: 21, ff. 14, 1 5 (1974); P. Royen, Alpine Fl.

STUDIES IN HYPERICUM

155

VANIMO

LORENGAU

to

<f

KAVIENG

Map 25 Sect. 26. 1. H. sewense *; 2. //. bifurcatum •; 4. //. macgregorii '

Atew Gwwea 3: 1472, f. 464 (1982). Type: New Guinea, Madang
Distr., Saidor subdistr., Finisterre Mts, Naho-Rawa Divide, Sewe,
Lake Naho, c. 2700 m, 13 November 1964 (fl & fr), Sayers NGF
21418 (BM!-holotype; L!-isotype).
Fig. 23A, Map 25.

Icon: Steenis, FL Males. I, 8: 20, f. 14 (1974).

Shrub c. 0.6 m tall, erect, with branches divaricate-ascending,
occasionally pseudo-dichotomous, mostly lateral. Stems 4-lined and
ancipitous when young, soon 2-lined, eventually terete, eglandular.
Leaves (7-)8-12 x 2-4 mm, lanceolate to narrowly elliptic,
concolorous, not glaucous, plane, spreading to ascending; apex

rounded, base narrowly cuneate; venation: 2-4 pairs of main lateral
veins branched distally, without tertiary reticulation; laminar glands
pale, linear to striiform towards base, sometimes flanked by row of
streaks or dots, punctiform towards apex and margin, inframarginal
glands dense, pale and (6-10) black. Inflorescence 1 -flowered,
without or rarely with one or two flowering branches in uppermost
axils, with numerous flowering branches from lower nodes; pedicels
4-9 mm long in fruit. Flowers 22-28 mm in diam., stellate; buds
narrowly ovoid, subacute. Sepals 5-7 x 1.5-2.5 mm, imbricate,
equal or subequal, lanceolate, rounded to subacute, entire; veins
5(3), outer branched; laminar glands pale, mostly linear to striiform,
punctiform towards apex and margin; inframarginal glands pale or
occasionally some black. Petals bright yellow, not tinged red, 10-14
x 5-6.5 mm, 2 x sepals, oblong-obovate, rounded, apiculus almost
absent; laminar glands pale linear, sometimes interrupted distally;
marginal glands absent or 1-3, black, sessile, on or near apiculus.
Stamens obscurely 3-fascicled, 1 5-20, longest (6.5-)7-8(-9) mm,
0.65-0.8 x petals; anther gland black. Ovary 2.5-3 x c. 1.5-2 mm,
narrowly ovoid, acute; styles 3, 2-4 mm long, 0.8-1.3 x ovary,
narrowly divergent; stigma scarcely capitate; placentae 3, intrusive
parietal. Capsule (5-)7-9 x (3-)4-5 mm, c. 1.1-1.4 x sepals,
ellipsoid to ovoid, with valves longitudinally vitiate. Seeds yellow-
brown, c. 0.8 mm long, scarcely carinate; testa densely
linear-foveolate.

Boggy tussock sedge-grassland; c. 2700 m.

New Guinea (Terr. New Guinea - Madang District).

NEW GUINEA. Madang Distr.: Saidor subdistr., Finisterre Mts, Naho-

156

N.K.B. ROBSON

Fig. 23 A. H. sewense: (a) habit; (b) leaf. B. H. papuanum: (c) habit; (d) leaf; (e) sepal; (f) petal; (g) capsule. C. H. saruwagedicum: (h) habit; (i) leaf (a,
c, h, x '/2; b, d, i x 2; e-g x 5). A. Sayers NGF 21418. B. Hoogland & Pullen 6012. C. Iserentant 9605.

STUDIES IN HYPER/CUM

Rawa Divide, Sewe, L. Naho, c. 2700 m, 1 3 November 1964 (fl & fr), Savers
NGF21418(BM, L).

H. sewense is in most respects the most primitive member of sect.
26. Humifusoideum, showing the greatest resemblance to the ances-
tral sect. 1 . Campy losporus. In that section, its affinities would seem
to be with//, lanceolatum subsp.angustifolium, which is endemic to
Reunion. Apart from its smaller size, it differs from that taxon in its
trimerous ovary, with relatively shorter divergent styles and parietal
placentation, its fewer 3-fasciculate stamens, and its usually rela-
tively broader leaves with more advanced venation and glandularity.
H. sewense is apparently a rare relict species, having been col-
lected only once.

2. Hypericum saruwagedicum Diels inBot. Jb. 62: 482 (1929); N.
Robson in Blumea 20: 257 ( 1 973), in Steenis, Fl. Males. 1, 8: 22,
ff. 16, 18e (1974); P. Royen, Alpine Fl. New Guinea 3: 1479, f.
466 (1982). Type: New Guinea, Morobe Distr., Saruwaged-
Gebirge, Bolan, 3400-3800 m, March-April 1913 (fl & fr),
Keysser s.n. (Bt-holotype; BM!-isotype?). The BM specimen is
labelled '3600-4000 m'.

Fig. 23C, Map 26.

H. macgregorii sensu Hoogland in Blumea, Suppl. 4: 231 (1958),

nonF. Mull. (1889).
H. macgregorii subsp. punctatum N. Robson in Blumea 20: 256

(1973), in Steenis, Fl. Males. 1, 8: 21 (1974); P. Royen, Alpine Fl.

New Guinea 3: 1478 (1982). Type: New Guinea, Irian Jaya, 6 km

NE of Lake Habbema, 3000 m, October 1938 (fl & fr), Brass

10660(A!-holotype).

Icon: P. Royen, Alpine Fl. New Guinea 3: 1480, f. 466 (1982).

Shrub or shrublet, 0.1-0.2 m tall, bushy, ericoid, with branches
strict, lateral, creeping and rooting at the base. Stems 4-lined when
young, soon 2-lined, eventually terete, eglandular. Leaves sessile;
lamina 2-9 x 0.5-3 mm, narrowly elliptic or lanceolate-elliptic to

157

linear, concolorous, not glaucous, incurved and proximally slightly
carinate, ± imbricate-appressed, rarely distally outcurving; apex
rounded, base narrowly cuneate; venation: c. 3 pairs of main lateral
veins, ± curved-parallel, unbranched except near apex and margin,
without noticeable tertiary reticulation; laminar glands pale, linear,
sometimes interrupted to punctiform near apex and margin, rarely
mostly striiform to punctiform; intramarginal glands spaced, pale
only or pale and black. Inflorescence 1 -flowered, with flowering
branches from scattered axils down stem; pedicels 2-4(-8) mm long
in fruit. Flowers 10-27 mm in diam., stellate or usually ± obconic;
buds narrowly ovoid-cylindric, subacute to rounded. Sepals 3.5-8 x
1-3 mm, imbricate or not, equal, ovate to lanceolate or elliptic to
narrowly oblong, subacute to rounded, entire; veins 7-9, forked and
distally branched; laminar glands pale, all or mostly linear or rarely
punctate; inframarginal to marginal glands pale only or pale and

Map 26 Sect. 26. 3. H. saruwagedicum 'forms': form 1 •; form 2 • ; form 3 D; form 4 A.

158

black. Petals dark to pale yellow, sometimes tinged red dorsally,
7-15 x 3-6 mm, c. 2 x sepals, obovate-oblong to oblanceolate-
spathulate, rounded, apiculus absent or almost so; laminar glands
pale, linear, sometimes interrupted distally; marginal glands absent
or rarely few, black, ± prominent. Stamens not obviously 3-fascicled,
( 1 3_)20-30, longest 4-7 (-9) mm, c. 0.5-0.75 x petals; anther gland
amber or black. Ovary 2-3 x c. 1 .5 mm, ovoid, acute; styles 3, 1 .5-
2.5 mm long, 0.5-0.65 x ovary, divergent; stigmas narrowly to
scarcely capitate; placentae 3, intrusive parietal. Capsule (5.5-)6-9
x 3.5-5 mm, c. 1.3-1.5 x sepals, ovoid, with valves longitudinally
vittate. Seeds orange-brown, 0.5-0.8 mm long, carinate; testa densely
linear-foveolate. 2n = 24 (Borgmann, 1964, as H. macgregorii).

Alpine grassland, open scrub or rocky slopes, usually in wetter

areas; mostly 2730-4300 m, but down to c. 1800 m in Milne Bay

District.

New Guinea (Irian Jaya - Mts Carstenz and Wilhelmina - to eastern

Papua - Mt Dayman).

IRIAN JAYA. 2 km E. of Wilhelmina-top, 3700 m, September 1938 (fl
& fr), Brass & Myer-Drees 101 19 (A, BM, BO, L); Mt Carstenz, IXA naar
Dajakweide, 3800-4300 m, November-Decenber 1936 (fl & fr), Wissel 133
(L); Lake Habbema, 3225 m camp, August 1938 (fl & fr), Brass 9043 (A,
BM. BO. K, L, LAE*).

TERR. NEW GUINEA Sepik Distr.: Telefomin subdistr., Sirius Mtn
and Sirius Plateau, 3000-3600 m, 23 April 1965 (fl), Craig 100 (LAE). E.
Highlands Distr.: Mt Wilhelm, vicinity of Mt Piunde, c. 3561 m, 1 August
1956 (fl), Womersley NFG 8874 (A, BM, BO, CANB*, K, L, LAE*, SING);
Kainantu subdistr., Mt Piora, 3150 m, 10 February 1963 (fl & fr), Henty &
Carlquist NFG 16561 (BO, CANB, K, L). Morobe Distr.: Salawaket Range,
Mamsin, c. 3820 m, 6 October 1964 (fl & fr), Hoogland 10002 (CANB*, K,
L, LAE*); Mt Amungwiwa, S. of Wau, 3420 m, November 1963 (fl & fr),
Womersley NGF 17979 (L, LAE*). Madang Distr.: Saidor subdistr., main
ridge of Finisterre Range, MtAbilala, c. 3390 m, 1 3 November 1964 (fl & fr),
Pullen 6070 (BM, L, LAE*).

PAPUA. S. Highlands Distr.: Mendi subdistr., Mt Giluwe, 3200 m, 25
December 1973 (fl), Croft et al. LAE 60678 (LAE). Central Distr.: Goilala
subdistr., Murray Pass, 2730 m, 25 July 1969 (fl & fr), Foreman NGF 45562
(K, L); Woitape subdistr., Avios, 1834 m, 25 August 1971 (fl), Millar 1208
(LAE). Milne Bay Distr.: Mt Dayman, Maneau Peak, 2780 m, 19 May 1953
(fl & fr), Brass 22250 (A, CANB*, K, L, LAE); Mt Suckling, 3330 m, 22
August 1965 (fl & fr), Gillison NGF 22384 (L, LAE*).

H. saruwagedicum is a variable species in which the variation falls
into four geographical but morphologically more or less intergrading
races. Although it is not desirable to give formal names to these
races, average members can be recognized by the following charac-
ters:

Variant 1 (Mt Wilhelm): Leaves large. Flowers large. Black
glands usually on leaves and sepals, not on petals or anthers. Shoot
apex outcurving.

Variant 2 (rest of Territory of New Guinea, west Papua): Leaves
medium. Flowers large to medium. Black glands on anthers only or
absent. Shoot apex erect. This variant is nearest morphologically and
geographically to 3. H. macgregorii.

Variant 3 (Irian Jaya): Leaves small. Flowers small. Black glands
on anthers and rarely leaves. Shoot apex erect.

In Flora Malesiana (Robson, 1974), H. saruwagedicum was dif-
ferentiated from H. macgregorii by its incurved (not flat) leaves with
laminar glands mostly linear (not mostly interrupted to punctate). In
addition, its ovary placentation is parietal (not axile); and it has black
glands on at least some part of the plant, whereas these are absent
from H. macgregorii An exception was found in one specimen from
Irian Jaya (L. Habbema), which has the black glands of H.
saruwagedicum but leaf characters otherwise similar to those of H.
macgregorii from east Papua. It was given subspecific rank under
the latter species, although geographically remote from it.

N.K.B. ROBSON

It now seems clear that the L. Habbema population is merely an
extreme form of Variant 3, the other specimens of which show a
trend towards it. H. macgregorii can therefore be regarded as
constituting another morphological trend from H. saruwagedicum
in which the black glands have disappeared and the leaves have
become smaller, flatter and relatively broader, with the glandular
pattern becoming more and more interrupted.

3. Hypericum macgregorii F. Mull, in Trans. R. Soc. Viet. 1(2): 2
( 1 889); Burkill in Kew Bull. 1899: 97 (1899); Lauterb. in Bot. Jb.
58: 4 (1922); Steenis in Bull. Jard. hot. Buitenz. Ill, 13: 219
(1934); N. Robson in Blumea 20: 255 (1973), in Steenis, Fl.
Males. 8: 21, if. 15, 18f (1974); P. Royen, Alpine Fl. New Guinea
3: 1476, t. 118 (1982). Type: New Guinea, Papua, Central
District, summit of Owen Stanley Range, 3900 m, 1 889 (fl & fr),
MacGregor s.n. (MEL!-holotype; BM!, BO!, SING!, WRSL!-
isotypes).

Map 25.

Icon: P. Royen, Alpine Fl. New Guinea 3: 1477, t. 1 18 (1982).

Shrub or shrublet 0. 1 5-1 .5 m tall, erect, bushy, with branches strict,
nearly always lateral, creeping and rooting at the base. Stems 2-lined
and ± ancipitous when young, eventually terete, eglandular. Leaves
sessile; lamina 4-9 x 1-3 mm, ovate-lanceolate to elliptic or nar-
rowly elliptic-oblong, concolorous, not glaucous, plane or ± incurved,
± imbricate-appressed to spreading; apex subacute to rounded,
margin plane or undulate, base narrowly cuneate; venation: 3(4-7)
pairs of main lateral veins, ± curved-parallel, not visibly branched
except near apex and margin, without tertiary reticulation; laminar
glands pale, linear towards base, interrupted or punctiform towards
apex and margin; intramarginal glands dense, pale. Inflorescence 1-
flowered, without or rarely with paired flowering branches in
uppermost axils, with scattered flowering branches from lower
nodes; pedicels 2-10(-20) mm in fruit. Flowers 20-25 mm in diam.,
stellate; buds narrowly ovoid, subacute. Sepals 3-6.5 x 0.75-3 mm,
not imbricate, equal, elliptic to linear-lanceolate, subacute to obtuse
or rarely rounded, entire; veins 7, outer ones branched; laminar
glands pale, all linear or some interrupted to punctiform towards
apex and margin; inframarginal glands pale. Petals dark to pale
yellow, sometimes tinged red dorsally, 7-15 x 3-6 mm, c. 2-2.5 x
sepals, obovate-elliptic, rounded, apiculus absent or almost so;
laminar glands pale, linear, sometimes interrupted distally, marginal
glands absent. Stamens not or obscurely 3(4?)-fascicled, c. 17-24,
longest 5-8 mm, c. 0.6-0.8 x petals; anther gland amber. Ovary 2 x
1 mm, ovoid, acute; styles 3(4), 1.5-2.5 mm long, 0.8-1.2 x ovary,
divergent; stigmas not capitate; placentae 3(4), axile. Capsule 6-8 x
3^ mm, c. 1 .3-2 x sepals, ovoid, with valves longitudinally vittate.
Seeds yellow-brown, c. 0.8 mm long, scarcely carinate; testa densely
linear-foveolate.

Open alpine rocks, wet slopes and roadsides, usually in shallow
soils; (1500-)2700^000 m.

New Guinea (Terr. New Guinea - Morobe District, Papua).

TERR. NEW GUINEA. Morobe Distr.: above village of Bakaia, c. 24
km SE of Garaina, c. 2700 m, 24 January 1964 (fl & fr). Hartley 12799
(CANB, L, LAE*).

STUDIES IN HYPER1CUM

159

PAPUA. Northern Distr.: Mt Scratchley, 2600 m, 1896 (fl & fr),
Giulianetti s.n. (K, MEL); Kokoda subdistr., E. side of L. Myolo, 2000 m, 22
July 1974 (fl & fr). Croft et al. LAE 61945 (LAE). Central Distr.: Goilala
subdistr., Mt Albert Edward, W. side, 3600 m, 21 June 1974 (fl & fr), Croft et
al. LAE 61389 (LAE); Port Moresby subdistr., Owen Stanley Range, head-
waters of Brown R., 1980m, 16July 1 969 (fl),/>a/>w™.y 798 (CANB*, LAE).
Milne Bay Distr.: Mt Dayman, Maneau [Maneao] Peak, 2780 m, 19 May
1953 (fl & fr). Brass 2225 1 (A, LAE*, MEL); Mt Maneao, 2250 m, 22 June
1956 (fl & fr), Cruttwell 746 (LAE).

Most specimens have a leaf index of 2.5-3.5 and fruiting pedicels 2-
6 mm long, but in the Mt Dayman (Milne Bay Distr.) population the
leaves are broader (1. i. = 2.2-3.1) and the fruiting pedicels longer
(5-10 mm). On the other hand, a population on the Mt Suckling
complex (Central/Milne Bay border) has larger leaves with puncti-
form glands and other characters intermediate between H.
macgregorli (or possibly H. saruwagedicum) and//, papuanum (e.g.
Milne Bay Distr., Raba Raba subdistr., S. end of Goe Dendiwa, 3430
m, 25 June 1972 (fl & fr), Stevens & Veldkamp LAE 54269 (A,
LAE). It seems likely that these are hybrids.

The complete absence of black glands is usually a good character
to separate H. macgregorii from H. saruwagedicum (q.v.), and they
apparently remain distinct where their areas overlap in eastern
Papua. In one population from Mt Albert Edward (Central Distr.) -
Brass 4168 (A, BM, BO, K, L, US), however, the gland pattern in
different plants varies from mainly linear to mainly punctiform, thus
forming a link between the two species.

4. HypericumbifurcatumN. Robsonin/?/w/m'a20: 256(1973), in
Steenis, FL Males. I, 8: 22, if. 15, 18d (1974); P. Royen, Alpine
Fl. New Guinea 3: 1474, f. 465 (1982). Type: New Guinea,
Morobe District, Huon Peninsula, Cromwell Mts, Mannasat, c.
2340 m, 9 August 1964 (fl & fr), Hoogland9542 (BM!-holotype;
CANB!, K!, L!-isotypes; LAE).

Map 25.

Icon: P. Royen, Alpine Fl. New Guinea 3: 1475, f. 465 (1982).

Shrub 0.3-1.5 m tall, with branches ± strict, pseudo-dichotomous

and lateral, sometimes also basal and rooting. Stems 2-4-lined when
young, eventually terete, eglandular. Leaves sessile; lamina 7-1 3(-
17) x 1.5-6 mm, narrowly ovate to narrowly elliptic-oblong,
concolorous, not glaucous, plane, ascending or appressed; apex
rounded, base cuneate to rounded; venation: 3-4 pairs of main
lateral veins, ± curved-parallel, scarcely branched except near apex
and margin, without noticeable tertiary reticulation; laminar glands
pale, linear to striiform, sometimes flanked by rows of dots, becom-
ing ± interrupted towards apex and margin; intramarginal glands
dense, pale. Inflorescence 1 -flowered, with paired strong flowering
branches in uppermost axils and often weaker ones in 1-2 axils
immediately below, repeated pseudo-dichotomies giving an effect
of bifurcations; pedicels 8-15 mm in fruit. Flowers 15-25 mm in
diam., stellate; buds narrowly ovoid, rounded. Sepals 4-6(-7) x 1 .5-
2 mm, imbricate, equal, ovate-lanceolate, subacute, entire; veins 7,
unbranched; laminar glands pale, all or mostly linear; inframarginal
glands pale or reddish. Petals bright yellow, orange- to red-tinged
dorsally, 9-14 x 3-5(-6) mm, 2.2-2.3 x sepals, obovate to
oblanceolate, rounded, apiculus absent or almost so; laminar glands
pale, linear, sometimes interrupted distally; marginal glands absent.
Stamens not or obscurely 3-fascicled, c. 25-30, longest (5-)6-8
mm, c. 0.75 x petals; anther gland black. Ovary 2-3 x 1.5-2 mm,
ovoid, acute; styles 3, 2-4 mm long, equalling or slightly longer than
ovary, divergent; stigmas narrowly capitate; placentae 3, parietal
except for axile extreme base. Capsule 6-9(-10) x 3.5-4.5 mm, c.
1 .3 x sepals, ± broadly to narrowly ovoid or ovoid-pyramidal, with
valves longitudinally vittate. Seeds yellow-brown, c. 0.7 mm long,
slightly carinate; testa densely linear-foveolate.

Wet to dry alpine grassland; 1 550-3000 m.

New Guinea (Terr. New Guinea - E. Highlands and Morobe Dis-
tricts).

TERR. NEW GUINEA. Morobe Distr.: Mt Salawaket [Sarawaket],
3000 m, 20 January 1963 (fl & fr), Hartley 1 1 166 (CANB, L, LAE*.); Busu
R. and vicinity, 1800-2400 m, 12 May 1937 (fl & fr), Clemens 6268 (A, B).
E. Highlands Distr.: Goroke subdistr., Marafunga, 3000 m, 29 August 1963
(fl & fr), Millar & van Royen NGF 1 5969 (CANB, L, SING); Chimbu Divide,
Daulo Road Camp, 2400 m, 6 November 1954 (fl), Womersley, Floyd &
McKee 6099 (A, K, LAE*).

The repeated pseudo-dichotomous inflorescence branching distin-
guishes //. bifurcatum from other members of sect. Humifusoideum
except sometimes H. papuanum, which has a different pattern of leaf
glands and often 4-5 styles. H. macgregorii rarely has one pseudo-
dichotomy, but the leaves are smaller, the anther gland amber and the
placentation axile. A collection of this species from a relatively low
altitude (1500 m) in the Owen Stanley Range (Central Papua) is
intermediate in this respect.

5. Hypericum papuanum Ridl. in Trans. Linn. Soc., Bot. 9: 19
(1916); Lauterb. in Bot. Jb. 58: 5 (1922); Steenis in Bull. Jard.
hot. Buitenz. 13: 219 (1934); N. Robson in Blumea 20: 257
(1973), in Steenis, Fl. Males. 8: 22, ff. 17, 18c (1974); P. Royen,
Alpine Fl. New Guinea 3: 1469, f. 463 (1982); Steenis in Blumea
28: 167 (1982). Types: Irian Jaya, Carstensz Peak, Camps 10-11,
1650-3300 m, 27 January 1913 (fl & fr), Kloss s.n. (BM!-
lectotype, selected here; K!-syntype); Carstensz Peak, Camps
1 1-12, 28 January 1913 (fl & fr), Kloss s.n. (BM!-syntype).

Fig. 23B, Map 27.

H.japonicum sensu Warb. in Bot. Jb. 16: 14 (1893).
H. macgregorii sensu Lauterb. in Nova Guinea 8: 843 (1912).
H. hellwigii Lauterb. in Bot. Jb. 58: 4 (1922); R. Keller in Engl. &
Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Steenis in Bull.

160

N.K.B. ROBSON

Map 27 Sect. 26: 5. H. papuanum •.

Jard. hot. Buitenz. Ill, 13: 219 (1934). Types: New Guinea, Terr.
New Guinea, Finisterre-Gebirge, c. 1 200 m, 1 5 October 1 888 (fl),
Hellwig 336 (Bt-lectotype, van Royen 1982 'holotype'; WRSL);
Irian Jaya, Treub-Gebirge, 2100-2300 m, 25 October 1909 (fl),
van Nouhys 1 (WRSLI-syntype); Irian Jaya, Hellwig-Gebirge,
2600 m, 2 December 1912 (fl), Pulle 594 (BO!-syntype); loc. cit.,
3 January 1913 (fl & fr), Pulle 890 (BO, L!, WRSL-syntypes).

H. habbemense A.C. Sm. in J. Arnold Arbor. 22: 343 (1941). Type:
New Guinea, Irian Jaya, 9 km NE of Lake Habbema, 2600-2650
m, October 1938 (fl & fr), Brass 10865 (A!-holotype; BM!, BO!,
L!, LAE-isotypes).

H. kunaianum Gilli in Annln naturh. Mus. Wien 83: 440 (1980).
Type: New Guinea, W. Highlands, Kuna Saw Mill bei Mt Hagen,
1900 m, Sommer 1970 (fl), Dosedla 76 (W!-holotype).

Icon: P. Royen, Alpine Fl. New Guinea 3: 1471, f. 463 (1982).

Shrub or shrublet 0.1-1.3(-2) m tall, densely or more usually
sparsely branched and ± spreading, with branches ascending, ± lax,
creeping and rooting at the base. Stems 2^-lined when young,
eventually terete, eglandular. Leaves sessile or subsessile; lamina 6-
25(-40) x 3- 1 7 mm, narrowly to broadly ovate or ovate-triangular to
elliptic or suborbicular, concolorous, not glaucous, plane, spreading
or subimbricate-ascending; apex subacute (or rarely acute) to
rounded, base rounded to cordate; venation: 4-5(6) pairs of main
lateral veins, ± curved-parallel or divergent, much branched to form
lax tertiary reticulation; laminar glands pale, linear towards base and
distally striiform to punctiform or wholly striiform to punctiform;
inframarginal glands dense, pale and/or black Inflorescence 1-
flowered, without or with flowering branches in uppermost axils, or
regularly dichasial or mixed dichasial/pseudo-dichotomous; pedicels
4-20 mm in fruit. Flowers 18-26 mm in diam., stellate; buds ovoid
to ellipsoid, subacute. Sepals 3-7(-8) x l-2.5(-3.5) mm, imbricate,
subequal to unequal, ovate to lanceolate or narrowly oblong or
sometimes broadly ovate and foliaceous, acute to rounded, entire;
veins 7-9, forking and distally branching; laminar glands pale,

linear; inframarginal glands black or absent. Petals bright yellow,
not tinged red, 9-15 x 4-9 mm, 2-3 x sepals, narrowly obovate-
elliptic, rounded, apiculus short or obsolete; laminar glands pale,
wholly linear or distally striiform; marginal glands absent or few to
numerous, black, often only in apiculus, not or scarcely prominent.
Stamens not obviously fascicled, ( 1 5-)25-40(-50), longest 6-9
mm, c. 0.75 x petals; anther gland amber or occasionally black.
Ovary (2-)2.5-3(-4) x 1.5-1.8 mm, narrowly or rarely broadly
ovoid, acute; styles 3(4-6), 2-3(^1) mm long, 0.75-1 x ovary,
divergent; stigmas narrowly to broadly capitate; placentae 3(4-6),
parietal. Capsule (5-)7-9(-10) x 3^4.5 mm, 1.3 x sepals, narrowly
or rarely broadly ovoid to ellipsoid, with valves longitudinally

STUDIES IN HYPERICUM

161

vitiate. Seeds yellow-brown to dark brown, 0.1-0.9 mm long, scarcely
carinate; testa densely linear-foveolate to linear-scalariform.

Wet or more rarely dry alpine grassland and bogs, screes, abandoned
cultivation; (1200-)! 600-3 800 m.

New Guinea (Irian Jaya (Mt Carstensz, Mt Hellwig, etc.) to Terr.
New Guinea (Madang District) and Papua (Milne Bay District)).

IRIAN JAYA. Central Distr.: Wissel Lake region, S. border of Lake
Paniai, foot of Mt Poti, February 1939 (fl & fr), Eyma 4531 (A, BM, BO, L,
LAE*); Bale R., 1 8 km NE of Lake Habbema, 2200 m, November 1938 (fl &
fr), Brass 1 1361 (A, BM, BO, K, L, LAE*). [Eastern Distr.?]: Mont. Hellwig,
2600 m, 3 January 1913 (fl), Pulle 890 (L).

TERR. NEW GUINEA. Sepik Distr.: Telefomin subdistr., Hindenburg
Range, Mt Kaban, 3120 m, 7 January 1965 (fl), Henty NGF 20655 (L). W.
Highlands Distr.: Wabag subdistr, N. slopes of Sugarloaf complex (along
Wapu R.), 2850 m, 1 2 July 1 960 (fl & fr), Hoogland & Schodde 7023 (A, BM,
CANB*, L, LAE*, PNH, SING, Z); Hagen subdistr., c. 0.4 km SE of Tomba,
c. 2400 m, 1 July 1957 (fl & fr), Sounders 649 (A, BM, CANB*, K, L, LAE*,
MEL, US). E. Highlands Distr.: Goroka subdistr., near Yontegi village,
between Dunantina R. and Karmanuntina R., c. 1850 m, 12 June 1956 (fl &
fr), Hoogland & Pullen 5310 (A, BM, CANB*, K, L, LAE*, MEL, PNH,
US); bottom of Mt Erimbari above Chuare, 2400 m, 26 January 1976 (fl),
Verdcourt & Johns 4930 (BM, K). Morobe Distr.: Sarawaket Range, SW
slope of Mt Enggom, along Zarun Creek, 2400 m, 24 February 1963 (fl & fr),
van Royen NGF 16143 (CANB, K, L, SING); Sattelberg, Lambanga, 1500-
1800 m, 14 September 1937 (fr), Clemens 7042a (A, B). Madang Distr.:
Saidor subdistr., Naho-Rawa Divide, Sewe, L. Naho, 2700 m, 13 November
1964 (fl & fr), Sayers NGF 21419 (BM, L).

PAPUA. Northern Distr.: Mt Simpson, 2880 m, 28 October 1947 (fl &
fr), Cruttwell 46 (K). S. Highlands Distr.: Mendi subdistr., W. slopes of Mt
Giluwe above Klareg, c. 2640 m, 25 October 1961 (fl & fr), Schodde 1971
(CANB, K, L, LAE, PNH); Turi subdistr., Mt Ambua, 3490 m, 13 October
1966 (fl & fr), Vmk 17433 (L, LAE). Milne Bay Distr.: Raba Raba subdistr.,
Mt Suckling complex, S. end of Goe Dendeniwa, 3430 m, 25 June 1972 (fl &
fr), Stevens & Veldkamp LAE 54269 (A, CANB*, LAE); Maneau Peak,
summit of Mt Dayman, 2780 m, 19 May 1953 (fl & fr). Brass 22244 (A,
CANB, L, LAE*).

H. papuanum is a very variable species in which the extreme forms,
although quite distinct in appearance, are linked by intermediates
with varying combinations of characters, so that the morphological
trends are not co-ordinated. These trends are:

(1) Leaves narrowly ovate and ± crowded, with laminar glands
mostly linear (in E., W. and S. Highlands mainly) to broadly ovate or
suborbicular, not crowded, with laminar glands all punctiform (con-
stant in Irian Jaya and east Papua - Northern and Milne Bay
Districts).

(2) Leaves, sepals, petals and anthers without black glands (mainly
eastern) to with black glands, forming a continuous inframarginal
row in the leaves (constant in Irian Jaya) and sepals and a continuous
marginal row in the petals (rare).

(3) Inflorescence 1 -flowered (mainly eastern) to regularly dichasial
(mainly western).

(4) Styles and placentae 3, with ovary and capsule narrowly ovoid
(mainly eastern) to styles and placentae 4-5 with ovary and capsule

Map 28 Sect. 26: 6. H. pulogense A; 7. H. beccarii: a. subsp. beccarii •; b. subsp. steenisii

162

N.K.B. ROBSON

± broadly ovoid (mainly western). The occurrence of 6 styles and 5
placentae reported by A.C. Smith (1941) was not confirmed on
examination of an isotype of H. habbemense. If correctly observed,
this character combination was no doubt teratological in origin.

(5) Habit dense with ± ascending branches (widespread) to lax
with ± spreading branches (Madang and Morobe Districts).

6. Hypericum pulogense Merr. in Philipp. J. Sci. 5, Bot.: 364
(1910), Enum. Philipp. ft. pi. 3: 75 1923); N. Robson in Blumea
20: 259 (1973), in Steenis, Fl. Males. I, 8: 24, f. 18a, b (1974).
Types: Philippine Is, Luzon, Benguet Prov., Mt Pulog, January
1909 (fl & fr), Curran, Merrill & Zschokke Philipp. For. Bur.
16097 (USMectotype, selected here; BM!, L!, PNHf, US!-
syntypes); Benguet Prov., Mt Pulog, May 1909 (fl), Merrill 6577
(K!, PNHt, US!-syntypes). Syntypes also include McGregor
8875 and 8880 (both PNH).

Map 28.

Icon: Steenis, Fl. Males. I, 8: 23, f. 18a, b (1974).

Shrublet or woody herb 0.2-0.4 m tall, diffuse?, with branches erect
or ascending to decumbent from slender branching rhizome. Stems
2(4-6)-lined or narrowly 2-winged when young, sometimes eventu-
ally terete, eglandular. Leaves sessile or very shortly petiolate;
lamina 8-12(-20) x 3-7 mm, ovate to elliptic or oblong, glaucous
beneath, plane (except margin?), spreading or ascending; apex
obtuse to rounded, margin recurved (at least when dry), base broadly
cuneate to rounded or subcordate; venation: c. 4 pairs of main lateral
veins, divergent-incurved, slightly branched to form obscure tertiary
reticulation; laminar glands pale, shortly striiform to punctiform,
subprominent beneath; intramarginal glands dense, pale. Inflores-
cence 1 -flowered, without or with flowering branches in uppermost
axils, or 3-10-flowered and regularly dichasial; pedicels 4-6 mm in
fruit Flowers 20-25(-30) mm in diam., stellate; buds narrowly
ovoid, subacute to obtuse. Sepals 4-6 x 1.8-2.4 mm, imbricate,
equal to subequal, lanceolate to oblong or elliptic-oblong, apiculate
to rounded, entire; veins 5, unbranched; laminar glands pale, linear
to punctiform; inframarginal glands pale, dense or rather sparse.
Petals bright yellow, not tinged red, 10-12(-14) x 4-6 mm, 2-2.5 x
sepals, narrowly oblong-ovate, rounded, with apiculus small, glan-
dular; laminar glands pale, numerous, basally linear, distally striiform
to punctiform; marginal glands few?, pale. Stamens not obviously
fascicled (5-fascicled/ufe Merrill), c. 30-60, longest c. 9 mm, c. 0.75
x petals; anther gland amber. Ovary 3-4 x 2 mm, ± broadly ovoid,
obtuse; styles 3, c. 5-5.5 mm long, c. 1 .5 x ovary, divergent; stigmas
not or scarcely capitate; placentae 3, axile. Capsule 5-8 x 4-5 mm,
narrowly to rather broadly ovoid, 1.3-1.5 x sepals, with valves
longitudinally and diagonally vittate. Seeds yellow-brown to dark
brown, 0.5-1 mm, not carinate; testa densely scalariform.

Summit grassland and open places in mossy forest; 2400-2800 m.
(see Jacobs, 1972).

Philippine Islands (northern Luzon).

PHILIPPINES. Luzon: Mountain Prov., Mt Pulog, 2550-2650 m, 30
January 1968 (fl & fr), Jacobs 7231 (K, L); Mt Tabayoc, 2400-2500 m, 15
February 1968 (fl), Jacobs 7437 (K, L).

H. pulogense, which is known from only Mts Pulog and Tabayoc, is
isolated morphologically as well as geographically. Having at first
related it (correctly) to the H. nagasawai complex of Taiwan but in
a distinct section, Pulogensia (Robson, 1973), I subsequently real-
ized that it has affinities with the New Guinea members of sect.
Humifusoideum, which it resembles in leaf and flower. The absence
of black glands in it occurs also in some populations of its apparently
nearest relative, 5. H. papuanum (q.v.), and the variation between
pseudo-dichotomous and dichasial inflorescence branching also
occurs in that species.

The Mt Tabayoc specimen has slender, more decumbent stems
than those from Mt Pulog, and in those characters it approaches 1 .H.
beccarii from Sumatra and Java.

6(i). Hypericum nagasawai Hayata in J. Coll. Sci. Univ. Tokyo
30(1): 38(1911), Icon, pi Formos. 1: 81, t. 18 (191 1); Makino
& Nemoto, Fl. Japan: 543 (1925), 2nd ed.: 749 (1931); Sasaki,
List pi. Formosa: 295 (1928); S. Susuki in Masamune, Short fl.
Formosa: 141 (1936);Y. Kimura in Bot. Mag. (Tokyo) 54: 82, f.l
(1940), in Nakai & Honda, Novafl. jap. 10: 225, ff. 75, 76.1
(1951); N. Robson in Li et al., Fl. Taiwan 2: 636, t. 431 (1976);
Li X.-W. in Fl. R. P. Sinicae 50(2): 70 (1990). Type: Taiwan,
Chaiyi, Mt. Morrison [Alishan] ad 13094 ped. [3928 m] ['top of
Mt. Yushan' on specimen], 2 November 1905 (fl), Nagasawa
754 (TI!-holotype).

Map 28a.

H. attenuatum sensu Hayata in J. Coll. Sci. Imp. Univ. Tokyo 25( 1 9):
59(1908).

H. randaiense Hayata in J. Coll. Sci. Imp. Univ. Tokyo 30( 1 ): 39
(1911), Icon, pl.formos. 1: 81, t. 17 (191 1)); Makino & Nemoto,
Fl. Japan: 295 (1925), 2nd ed.: 752 (1931); Sasaki, List pi.
Formosa: 295 (1928); S. Susuki in Masamune, Short fl. Formosa:
141 ( 1936); Y. Kimura in Bot. Mag. (Tokyo) 54: 84, f. 2 (1940), in
Nakai & Honda, Nova fl. jap. 10: 226, f. 76.3 (1951). Type:
Taiwan, Nantou, Randaisan [Luantashan], 8 August 1908 (fl),
Kawakami & Hayata s.n. (TI!- holotype).

H. nagasawai var. typicum Y. Kimura in Bot. Mag. (Tokyo) 54: 82
(1940), in Nakai & Honda, Novafl. jap. 10: 225 (1951). Type as
for H. nagasawai.

H. nagasawai var. nigrumY. Kimura in Bot. Mag. (Tokyo) 54: 82, f.
Ic (1940), in Nakai & Honda, Novafl. jap. 10: 225 (1951). Type:
Taiwan, Nantou, Prov. Taityu, inter Noko et Boarun, 1 6 June 1930
(fl), Kudo & Mori 2308A (TAI! -holotype).

H. taiwanianumY. Kimura in Bot. Mag. (Tokyo) 54: 84, f. 3 (1940),
in Nakai & Honda, Novafl. jap. 10: 227, f. 76.6 (1951). Type:
Taiwan, Taichung, Mt. Silvia [Tugitakayama], 3000 m, 13 July
1924 (fl), S. Ohasi in Herb. Simada 1214 (TI!- holotype).

H. hayatae Y. Kimura in Bot. Mag. (Tokyo) 54: 85, f. 5 (1940), in
Nakai & Honda, Nova fl. jap. 10: 228, f. 76.4 (1951). Type:
Taiwan, Taichung, Hakku-taisan [Paikoutashan], 9 August 1908
(fl), Mori s.n. (TI! -holotype).

H. suzukianum Y. Kimura in Bot. Mag. (Tokyo) 54: 86, f. 6 ( 1 940), in
Nakai & Honda, Novafl. jap. 10: 228, f. 76.2 (1951). Type:
Taiwan, Taichung, Prov. Taityu, Kunigigaoko [Kunugioko], 27
July 1936 (fl), Suzuki in HTU 1 18999 (TAI).

H. taiwanianum var. taiwanianum Y. Kimura in Nakai & Honda,
Novafl. jap. 10: 227 (1951), autonym.

H. taiwanianum var. ohwiiY. Kimura in Nakai & Honda, Novafl.

STUDIES IN HYPER/CUM

163

Map 28a Sect. 26: 6(i). H. nagasawai •.

jap. 10: 228 ( 1 95 1 ). Type: Taiwan, Ilan, Painan-ambu, n.d. (fl), S.
Ohwi s.n. (TI!- holotype).

Icones: Hayata, Icon, pl.formos. 1: t. 18 (191 1); N. Robson in Li et
al., Fl. Taiwan 2: t. 431 (1976).

Perennial herb or deciduous shrublet, 0.05-0.35 m tall, suberect to
ascending from creeping branching rooting base, with stems solitary
or ± caespitose, unbranched or ± branched above, wiry. Stems 2(4)-
lined, eglandular; internodes c. 5-15 mm, shorter than leaves.
Leaves sessile or subsessile; lamina (3-)8-25 x 3-12 mm, ovate or
oblong to elliptic or oblanceolate or linear, glaucous or sometimes
minutely papillose beneath, chartaceous to subcoriaceous; apex
acute to rounded, margin entire, recurved, base cuneate to angustate;
venation: 3-4 pairs of main laterals from lower third of midrib,
sometimes forming ± marked submarginal vein, tertiary venation
lax or obscure; laminar glands pale, somewhat elongate to puncti-
form, rather sparse, prominent above; intramarginal glands black,
dense or irregular. Inflorescence 1-1 1 -flowered from 1-2 nodes,
sometimes with flowering branches from 1-2 nodes below, the
whole subcorymbiform; pedicels 4-8 mm; bracts and bracteoles 2-
6 mm long, lanceolate to linear, entire. Flowers 15-30 mm in diam.,
± stellate; buds ovoid to ellipsoid, obtuse. Sepals 5, equal, 3-7.5 x
0.8-2.5 mm, in bud and fruit, ovate-lanceolate to ± narrowly oblong,
obtuse to acute, entire or rarely with 1-2 glandular cilia; veins 5,

outwardly branched; laminar glands pale or rarely black, striiform to
punctiform or rarely linear; marginal glands black and sometimes
pale, regular or irregular, immersed or rarely on cilia, or absent.
Petals 5, bright yellow, not tinged red in bud, 8-1 7 x 4-7 mm, 2-2.5
x sepals, obovate or oblong-obovate to oblanceolate, usually entire,
laminar glands pale and rarely black, linear to punctiform, or rarely
absent, marginal glands black, sessile or occasionally on cilia, distal
and few or subapical and solitary. Stamens 40-80, not or obscurely
fascicled, longest 4.5-8 mm, 0.5-0.8 x petals; anther gland black.
Ovary 3-locular, 2-2.5 x 1-1.5 mm, narrowly ovoid to ellipsoid-
ovoid; styles 3, 3.5-7 mm, 1.3-3 x ovary, ± spreading from near
base; stigmas narrowly or scarcely capitate. Capsule (5-)6-7 x 3.5-
5 mm, c. 1.5 x sepals, narrowly to broadly ovoid; valves narrowly
longitudinally vittate. Seeds dark brown, c. 1 mm, not or scarcely
carinate, apiculate; testa finely scalariform-reticulate to linear-fo-
veolate. 2n = 36 (n = 18, Hsu, 1968).

Stony or rocky slopes, roadsides and open areas of conifer forests
and subalpine woodland; 2300-3997 m.

Taiwan (central mountains).

TAIWAN. Ilan: Chi-li-tin to Nanfu-shan-chuang, 21 August 1969 (fl),
Hsu 5930 (TAI); in Ml. Nanko-taisan, July 1933 (fl), Ohwi 4095 (K).
Hsinchu: Mt. Isawa, 18 July 1932 (fl), Sasaki in HTU 077131 (TAI); Mt.
Taiha, on top, 5 July 1934 (fl), Suzuki in HTU 077087 (TAI). Taichung: Mt.
Tugitaka, 19 August 1930 (fl), Onutna 12 (TAI); Souyuan-akou (Piyanan-
ambu), S. slope of Mt. Layeh-wei-shan, 200-2300 m, 10 July 1963 (fl),
Shimizu & Chuang 20142 (E). Nantou: southern flank ofYushan, 3860 m, 30
October 1992 (fl), Kirkham & Flanagan ETOT170 (K); Ten-tzu to Nen-kao,
12 August 1971 (fl), Huang, Hsieh & Kao 5825 (TAI). Chiayi: Arisan
[Alishan], 2500 m, July 1914 (fl), Faurie 536 (BM); Pai-yunn Hostel to top
of Mt. Morrison [Alishan], 3550-3997 m, 6 September 1969 (fl), Hsu 6283
(TAI). Hualien: Ko-Nan-Kuan, 16 July 1966 (fl), Chuang & faze 4268 (TAI,
US); fromTayn Lin to pass at Ko-nan Kuang, 3000 m, 8 August 1966 (fl), van
Steenis 20700 (L).

CULTIVATED. England: Kew, seed ex Formosa, Yashiroda 88, 16 Au-
gust 1 934 (fl), Anon. (K). Ireland: Co. Meath, Kells, I.F.S., cult. Marquess of
Headfort, seed coll. Formosa ex Yashiroda 88, 13 August \936(ft),Anon. (K).

H. nagasawai is quite closely related to//, pulogense, differing from
it inter alia by the more slender habit, the frequently narrower leaves
and the black glands on the anthers and elsewhere. The variation
from the mainly northern broad-leaved form with sepals and styles
about 1 .3 times as long as the ovary (H. nagasawai) to the southern
narrow-leaved form with acute sepals and styles 2 or more times as
long as the ovary (H. randaiense) appears to be continuous; and
indeed the trends in leaf-form, sepal-shape and style-length are only
partially correlated. It is not possible, therefore, to recognize H.
randaiense as a distinct species. Similarly, plants described as H.
suzukianum and//, hayatae, respectively, are more extreme forms of
trends within H. nagasawai; and the occasional forms with black-
glandular-ciliate sepal margins (H. taiwanianum) are linked to the
more typical forms by specimens with one or two glandular cilia on
each sepal margin.

6(ii). H. nokoense Ohwi InActa Phytotax. Geobot. 6: 48 ( 1937);Y.
Kimura in Bot. Mag. (Tokyo) 54: 85, f. 4 (1940), in Nakai &
Honda, Nova fl. jap. 10: 226, f. 76.5 (1951); N. Robson in Li
et al., Fl. Taiwan 2: 639 (1976); Li, X.-W. in Fl. R. P. Sinicae
50(2): 69 (1990). Type: Taiwan, Hualien, Mt. Nokogoe, in
Karenko, June 1933 (fl), Ohwi 2951 (KYO-holotype); loc.
cit., June 1933 (fl), Ohwi 2958 (K!, KYO, Tl-paratype).

Map 28b.

Icon: Y. Kimura in Nakai & Honda, Nova fl. jap. 10: 226, f. 76.5
(1951).

164

N.K.B. ROBSON

Map 28b Sect. 26: 6(ii). H. nokoense •

Perennial herb 0.05-0.1 m tall (sometimes longer and straggling),
suberect to ascending from creeping, branching and rooting base,
with stems ± caespitose, sometimes mat-forming, unbranched or ±
branched above. Stems 2-4-lined or sometimes becoming almost
terete, eglandular; internodes 2.4-6 mm, usually longer than leaves.
Leaves sessile or with petiole up to 1 mm; lamina 4-12x1 .5-6 mm,
ovate (below inflorescence) to elliptic or narrowly oblong or obovate,
markedly glaucous and sometimes minutely papillose beneath,
subcoriaceous; apex rounded, margin entire, base cuneate-angustate;
venation: 2-3 pairs of main laterals from lower 1/3 to 2/5 of midrib,
tertiary reticulation obscure; laminar glands pale and sometimes
black, punctiform, dense to sparse; intramarginal glands black,
dense. Inflorescence l-5(-7)-flowered, from apical node, without
lower branches, subcorymbose; pedicels 1-1.5 mm; bracts and
bracteoles 3-3.5 mm long, entire or with prominent marginal glands.
Flowers 12-18 mm in diam., ± stellate; buds ovoid to ellipsoid,
acute. Sepals 5, equal, erect in bud and fruit, 3-5 x 0.8-1.2 mm,
lanceolate to narrowly oblong, acute to subacuminate, entire; veins
3-5, unbranched; laminar glands pale and black, linear to striiform;
marginal glands black, irregular, submarginal. Petals 5, bright yel-
low, not tinged red in bud, 7-11 x 2-3(^.5) mm, c. 3 x sepals,
obovate or oblong-lanceolate to lanceolate, laminar glands pale and
usually black, linear to punctiform, marginal glands absent. Stamens
c. 40-43, obscurely '3'-fascicled, longest 5-8 mm, c. 0.7 x petals;

anther gland black. Ovary 3-locular, c. 2 x 1 mm, ovoid?; styles 3,
free, 4-6 mm, 2.5-3 x ovary, spreading from base; stigmas not
enlarged. Capsule 6-7 x c. 4 mm, c. 1 .5-2 x sepals, narrowly ovoid;
valves longitudinally vitiate. Seeds not seen.

Mountain slopes; 1800-1900 m.

Taiwan (Nantou, Hualien).

TAIWAN. Nantou: Sakahen to Kiraikei, 2 1 August 1929 (fl), Sasaki in
HTU 077267 (TAI). Hualien: Mt. Luan-shan, 17 October 1967 (fl), Huang
4218 (TAI); Mt. Chin-shuei-shan, 21 June 1941 (fl), Nakamura 5355 (TAI).

No single character separates H. nokoense from H. nagasawai (from
which it appears to have been derived by reduction); but the combi-
nation of small and relatively broad leaves, acute sepals usually with
black laminar glands, and styles more than twice as long as the ovary
appears to distinguish it. The leaf glands are not wholly black (pace
Ohwi and Kimura).

7. Hypericum beccarii N. Robson in Blumea 20: 260 (1973), in
Steenis, FL Males. I, 8: 25 (1974). Type: Sumatra, Barat,
Tadangsche Bovenland', G. Singgalan [Singalang] June-July
1878 (1. fl & fr), Beccari 337 (BM!-holotype; K!, L!, MEL!-
isotypes).

Map 28.

H.japonicum var. pinnatinervium Bakh. f. in Backer & Backh. v. d.
Brink, FL Java 1: 382 (1963), nom. illegit. descr. angl. (Art.
36.1).

Perennial (or sometimes annual?) herb c. 0.02-0.45 m long, branch-
ing irregularly, wiry, with branches from near base, weak, decumbent
or ascending (erect fide Bakh. f.), creeping and rooting. Stems 4-6-
lined when young, often becoming 2-lined, eglandular. Leaves with
petiole 0.2-1.5 mm long; lamina 2.5-10.5 x 0.5-6 mm, broadly
oblong or elliptic-oblong to oblanceolate or linear, glaucous be-
neath, plane (sometimes except margin), spreading; apex subacute
or apiculate to rounded, margin not or slightly recurved, base
rounded to cuneate; venation: 3(4) pairs of main lateral veins, ±
parallel-incurved, branched to form tertiary reticulation dense and
conspicuous near apex and margin; laminar glands pale, irregularly
punctiform; intramarginal to marginal glands irregular, pale and/or
black. Inflorescence 1 -flowered, with flowering branches in one or
both uppermost axils; pedicels (2-)5-17 mm in fruit. Flowers c. 1-
10 mm in diam., stellate; buds narrowly ovoid, obtuse. Sepals 2.5-5
x 0.6-1 .4 mm, imbricate or not, unequal to equal, elliptic-oblong to
linear, rounded to apiculate or subacute, entire to irregularly glandu-
lar-ciliate; veins 5, branching and reticulating; laminar glands pale,
shortly striiform to punctiform; inframarginal to marginal glands
black, spaced. Petals yellow, not tinged red, 3-7 x 2-2.5 mm, 1.2-
1.4 x sepals, oblong-oblanceolate, rounded, with apiculus glandular
or glandular-ciliate; laminar glands pale or black, few, near apex,
punctiform, or absent; marginal glands black, 1-2, subsessile or on
cilia, sometimes continuing as a row of inframarginal to submar-
ginal glands. Stamens clearly (?) 3-fascicled, c. 15-22, longest
2.5-5.5 mm, c. 0.75-0.85 x petals; anther gland black. Ovary 1.5 x
c. 1 mm, ± narrowly ovoid, acute; styles 3, c. 1 .5 mm long, placentae
3, axile. Capsule 3-5.5 x 2-3.5 mm, ± narrowly ovoid, 1.3-1.2 x
sepals, with valves longitudinally vittate or almost smooth. Seeds
reddish brown, 0.7-0.9 mm, not carinate; testa densely and shallowly
linear-reticulate.

Damp places in open vegetation; 1800-3000 m.

Sumatra (N., W.-central), Java (W.).

H. beccarii is more closely related to 5. H. papuanum than to 6. H.

STUDIES IN HYPER1CUM

pulogense, in particular to the small, more diffuse form from Irian
Jaya (which includes the type). The most primitive characters occur
in the central Sumatran populations on G. Talamau and G. Singalang,
those on Java (on G. Papandajan only) and northern Sumatra being
dwarfer with a cuneate leaf base. The northern Sumatran population,
which also differs in its narrower leaves and almost smooth capsules,
is worthy of distinction as a subspecies.

7a. Hypericum beccarii subsp. beccarii

165

Leaves with petiole 0.2-1 mm long; lamina 4-1 0 x 2-6 mm, broadly
oblong or elliptic-oblong to narrowly obovate; apex rounded-
apiculate to rounded, base rounded to cuneate; laminar glands ±
prominent. Sepals 3.5-5 x 1-3 mm, ± broadly imbricate. Petals 5-
7 mm long; laminar glands (when present) black. Stamens c. 20-22,
longest 4.5-5.5 mm. Capsule 4-5.5 x 2.5-3.5 mm, densely and
prominently vittate.

1 500-3000 m.

Sumatra (W. -central, on GG. Singalang, Talang, Talamau and
Kerintji), Java (G. Papandajan).

SUMATRA. Jambi: Talang, Taloe, 1500 m, 15 June 1917 (fl & fr),
Bunnemeijer 1050a (BO); Talamau, NW slope, 2700 m, 25 May 1917 (fl &
fr), Bunnemeijer 843 (BO). Barat: G. Koerintji, 2400 m, 10 May 1920 (fl),
Bunnemeijer 10398 (BO); G. Indrapura, 3000 m, 11 January 1914 (fl),
Matthew s.n. (K).

JAVA. Priangan: G. Papandajan, c. 2450 m, 21 January 1930 (fl & fr),
van Steenis 4080 (L).

Specimens with broad leaves and large flowers (from G. Talamau and
G. Singalang) approach H. papuanum most closely. The Javan and G.
Indrapura populations are dwarfer with smaller, cuneate-base leaves,
relatively shorter pedicels and sepals more often with glandular cilia,
but they cannot otherwise be separated from those further north.

7b. Hypericum beccarii subsp. steenisii N. Robson in Blumea 20:
261 (1973), in Steenis, Fl. Males. 1, 8: 25 (1974). Type: Sumatra,
Gaju and Alas Lands, Mt Losir, bivouac 4 to 5, 2700-2800 m, 3 1
January 1937 (fl & fr), van Steenis 8514 (L!-holotype; A!, K!-
isotypes).

Leaves with petiole 0.5-1.5 mm; lamina 2.5-10.5 x 0.5^ mm,
oblanceolate to linear; apex rounded to subacute, base cuneate;
laminar glands not prominent. Sepals 2.5-5 x 0.6-1.4 mm, not
imbricate. Petals 3-6 mm long; laminar glands (when present) pale.
Stamens c. 1 5-20, longest 2.5-^.5 mm. Capsule 3-4.5 x 2-3 mm,
sparsely and obscurely vittate (almost smooth).

Along streamlets in open vegetation; 2700-3314 m.

Sumatra (N.).

SUMATRA. Aceh: Gunung Leuser National Reserve, between and
below G. Leuser West top and Middle top, 3200 m, 10 April 1975 (fl), de
Wilde & deWilde-Duyfies 1 6325 (K); Gajo Lands, G. Kemiri, E. side, bivouac
2 near top, 3100-3314 m, 7 March 1937 (fl), van Steenis 9599 (L).

8. Hypericum natalense J.M. Wood & M.S. Evans in J. Bot. Land.
35: 487 (1897); R. Keller in Engl. & Prantl, Nat. Pflanzenfam.
2nd ed. 21: 177 (1925); Burtt Davy, Man. pi Transvaal 1: 25
(1926); Bredell in Bothalia 3: 579 & map (1939); N. Robson in
Kew Bull. 12: 440, map 2 ( 1 958); Killick & Robson in FL southn
Afr. 22: 19 (1976). Type: Natal, Estcourt District, near bank of
Mooi river, 1200-1500 m, 26 October 1888 (fl & fr), Wood 4034
(NH-holotype; BM!, BOL, G, K!, Z!-isotypes; PRE-photograph).

Fig. 24A, Map 29.

H. woodii R. Keller in Bot. Jb. 58: 193 (1923), in Engl. & Prantl,
Nat. Pflanzenfam. 2nded. 21: 179 (1925). Type: Natal, near bank
of Mooi river, 1 200 m, 26 October 1 888 (fl & fr), Wood NH 788
(Bt-holotype). Keller's type is from the same collection as that of
Wood & Evans.

H. natalense van petiolatum Bredell in Bothalia 3: 580 ( 1 939). Type:
Natal, Camperdown District, no precise locality, Franks NH
12968 (NH-holotype; PRE-photograph).

Perennial woody herb 0.2-0.45 m tall, fasciculate, with branches 1-
many from underground rootstock, erect, much branched. Stems
4-lined and ancipitous when young, soon terete, eglandular. Leaves
sessile or rarely to 0.5 mm petiolate; lamina 9-20 x 5-12 mm,
broadly elliptic to obovate, paler but not glaucous beneath, plane,
spreading; apex obtuse to rounded, base cuneate to rounded; vena-
tion: 1-3 pairs of main lateral veins, much branched to form fairly

166

N.K.B. ROBSON

Map 29 Sect. 26. 8. H. natalense • specimens, D records; 9. H. wilmsii A specimens, A records; 10. H. peplidifolium • specimens, O records.

conspicuous tertiary reticulation; laminar glands pale, punctiform;
intramarginal glands spaced, black. Inflorescence 1 -flowered, with
(usually) paired flowering branches from 1-2 or more (up to 10)
nodes below; pedicels 4-10 mm in fruit. Flowers c. 10-14 mm in
diam., stellate; buds ellipsoid, obtuse. Sepals 4-8 x 1.5-4 mm,
imbricate, unequal, elliptic to obovate or spathulate, rounded-
subapiculate to rounded, entire; veins 5, branched and reticulating;
laminar glands pale, dense, shortly striiform to punctiform; infra-
marginal glands few, mostly subapical, black. Petals yellow, not
tinged red, 6.4-7 x 1.7-2.5 mm, c. 0.9-1.5 x sepals, elliptic to
oblong or spathulate, rounded, apiculus subterminal, obscure to
obsolete; laminar glands few, pale or absent; marginal glands absent

or few, black, near apex. Stamens irregularly 3-fascicled or not
fascicled, c. 30, longest 3-5 mm, c. 0.5-0.7 x petals; anther gland
black. Ovary 2.5-3 x 2 mm, ovoid-cylindric, obtuse; styles 3-^-(5),
2-2.5 mm long, 0.8-0.9 x ovary; stigmas narrowly capitate; placen-
tae axile. Capsule 5-6 x 3 mm, ovoid-cylindric, 1.25-1.35 x sepals,
with valves longitudinally vittate. Seeds yellow-brown, 0.7-0.9 mm,
not carinate; testa scalariform-reticulate.

Damp places in grassland; 300-1650 m.

Eastern Transvaal, Swaziland, Natal (Midlands), eastern Cape Prov-
ince.

STUDIES IN HYPERICUM

167

Fig. 24 A. H. natalense: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) capsule. B. H. wilmsii: (f) habit; (g) capsule; (h) sepal. C. H. peplidifolium: (i) habit; (j)
sepal;(k) petal; (1) 'berry' (a, f, i x Vr, b x 2; e, h, 1 x 4; c, d. g, j, k x 5). A. Medley Wood 625 1 . B Exell, Mendon^a & Wild 1 24. C. Newman & Whitmore 19.

168

N.K.B. ROBSON

TRANSVAAL. Pilgrim's Rest Distr., Mac Mac, pre-July 1884 (fl),
Mudd s.n. (K).

SWAZILAND. Mbabane, Poliniane R., Compton 26489 (NBG*).8

NATAL. Pietermaritzburg Distr., mountain top before Hela Hela, 2
November 1969 (1. fl), Strey 9221 (K); Pietermaritzburg Distr., Howick
[Falls], 300 m. 1893 (fl), Junod 217 (Z); Umvoti Distr., Greytown, 968 m,
October-November 1931 (fl), Wylie s.n. (K); Houtbosh, 1875-1880 (fl),
Rehmann 6306 (Z); Estcourt Distr. (see type); Mpendhle Distr., Litani area,
Elandshoek Valley, 1650 m, 26 December 1984 (fl & fr), Milliard & Burn
1804(K).

CAPE PROVINCE. Griqualand, Mt Fletcher Distr., Mt Fletcher, No-
vember 1913 (fl), Jacottet & Jacottet 570 (Z);Tsolo Distr., Payne 14 (GRA*);
Umtata Distr., Baziya, 600 m, pre 1885 (fl & fr), Baur 582pp. (K); Kentani
Distr., October, Pegler 1 17 (GRA*).

H. natalense is related to 4. H. bifurcation, differing from it in being
less woody with stems more branched basally and laterally, so that
there are lateral branches bearing repeated pseudo-dichotomies
from usually several lower stem nodes. In addition, the leaves are
thinner and spreading and the leaf laminar glands are punctiform,
not linear. Despite the wide geographical separation, however, there
can be no doubt of the close relationship of these two species.

9. Hypericum wilmsii R. Keller in Bull. Herb. Boissier II, 8: 179
(1908) in clav., in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21:
1 8 1 ( 1925); Bredell in Bothalia 3: 579 (1939); N. Robson in Kew
Bull. 12: 440, map 2 (1958), in Exell & Wild, Fl. Zamb. 1: 383
(1961); Killick & Robson in J. Ross, Fl. southn Afr. 22: 20
(1976). Type: Transvaal, Lydenburg District, bei der Stadt
Lydenburg, January 1888 (fr), Wilms 136 (W!-holotype; PRE!,
fragment).

Fig. 24B, Map 29.

H. rupestre sensu H. Perrier inArchs Bot. Bull. mens. 1: 9, 1 1 ( 1 927)
pro parte quoad descr., nee Jaub. & Spach (1 842) nee Bojer ex R.
Keller (1925).

H. nigropunctatum Norl. in Bot. Notiser 1934: 103, t. 8 (1934).
Type: Zimbabwe, Inyanga, c. 3 km occidentum versus a monte
Inyangani, c. 1900 m, 8 December 1930 (fl & fr), Fries, Norlindh
& Weimarck 3634 (LD!-holotype; BM!, PRE!, SRGH!-isotypes;
K!-photograph).

H. bojerianum sensu H. Perrier in Not. Syst. Paris 13: 269 (1948), in
Mem. Inst. sclent. Madagascar 1: 1 13 (1949), in Humbert, Fl.
Madag., Hyperic.: 4, f. I 14-17 (1951), pro parte omnes excl.
typum. See Robson (1958: 440).

H. aethiopicum sensu Jacot Guill., Fl. Lesotho: 211 (1971) (as H.
aethiopicum subsp. sonderi) pro parte?, see p. 169.

Icon: Humbert, Fl. Madag., Hyperic.: 5, f. I 14-17 (1951).

Perennial herb, 0.06-0.2 m tall, spreading or straggling, with
branches ± numerous from taproot, decumbent or ascending,
branched, sometimes rooting. Stems 2-lined and ± ancipitous above,
otherwise terete, eglandular or densely black-gland-dotted. Leaves
with petiole to 2 mm long or rarely almost sessile; lamina (3-)6-10
x 2-7 mm, ovate or elliptic to obovate or suborbicular, plane
spreading; apex rounded, margin slightly recurved, base rounded to
cuneate; venation: 2-4 pairs of main lateral veins, branching to form
dense inconspicuous tertiary reticulation; laminar glands pale, punc-
tiform; inframarginal glands dense, all or mostly black. Inflo re. scence
1 -flowered, with paired flowering branches from uppermost and
sometimes next lower nodes; pedicels 3-10 mm, erect in fruit.
Flowers 9-12 mm in diam.; buds ellipsoid, obtuse. Sepals 4-7 x 1-
2 mm, imbricate, unequal, all oblong or outer ones obovate, obtuse

8 An asterisk (*) after records of South African specimens of sect. Humifusoideum
indicates that they have been seen by Killick but not by me.

or apiculate to rounded, entire; veins 3, branched and reticulating;
laminar glands pale, punctiform; inframarginal glands numerous to
few, black or rarely absent. Petals bright to primrose yellow, occa-
sionally tinged red, 5-9 x 2-3.5 mm, c. 1.3 x sepals, oblong to
obovate, apiculus subterminal, obscure; laminar glands pale or
rarely some black streaks; marginal glands sparse, black, near apex.
Stamens irregularly 3-4-fascicled or not fascicled, 1 8-30, longest c.
4-6 mm, c. 0.7 x petals; anther gland black. Ovary 2.5^4- x 2 mm,
obtuse; styles 3^4, ( 1 .5-)2-2.5 mm long, c. 0.65 x ovary, divergent;
stigmas narrowly capitate; placentae 3-4, axile. Capsule 5-6 x 3-5
mm, ellipsoid, 0.8-0.9 x sepals, erect, with valves longitudinally
vittate. Seeds yellowish brown, 1-1 .5 mm, not carinate; testa linear-
foveolate.

Damp places and on mountain slopes; 1200-2000 m (Zimbabwe),
1200-1500 m (South Africa), 1200-2500 m (Madagascar).

Zimbabwe (E.), Transvaal, Orange Free State, Lesotho?, eastern
Cape Province, Madagascar.

ZIMBABWE. Eastern: Umtali Distr., Emgwa, 1980 m, 2 February 1955
(fl & fr), Exell, Mendonfa & Wild 124 (BM, SRGH); Umtali Distr., Himala-
yas, Banti North, 1950 m, 4 March 1964 (fr), Wild 45 19 (K, SRGH); Inyanga
Distr., N. side of Pungwe Gorge, 3 February 1957 (fl & fr), SeagriefCAH
1 026 (K, SRGH*).

TRANSVAAL. Lydenburg Distr. (see type of H. wilmsii).

ORANGE FREE STATE. Bloemfontein, Thaba Mchu, Roberts 2350
(PRE*).

LESOTHO (see below). Mafeteng Distr.: Likiele, Mt Ha-moya-pela, 6
January 1916, Dieterlen 1 222 (PRE*); Catai R., Ha-Ma-Khonofane, Dieterlen
1293 (PRE*).

CAPE PROVINCE. Herschel, Majubanek, near Sterkspruit, December,
Hepburn 92 (GRA*). Aliwal North: Doctor's Drift, 1233 m, 8 December
1933, Gerstner 137 (PRE p.p.*); Elandshoek, c. 1356 m, October, Bolus 153
(PRE*, SAM*). Queenstown: no precise locality, 1200-1500 m, November-
December (fl), Galpin 1629 (PRE*). Somerset East: Cradock Distr., Mt
Zebra Park, 13 November 1953, Brynard 291 (PRE*); Somerset East, 6
March 1866 (fl & fr), Bolus 319 (K, TCD*).

MADAGASCAR. Centre (mountains to S. of Imerina): Ankaratra au N.
de Antsirabe, Perrier 14626 (P*); Betafo, a 1'W. de Antsirabe, Perrier 3467
(P*); S. Betsileo, environs d'Ambalavao, Perrier 14625 (P*); massif
d' Andringitra, au S. d'Ambalavao, Perrier 3663 (P*).

H. wilmsii is intermediate in form and distribution between 8. H.
natalense and 1 0. H. peplidifolium, occurring to the west and north
of the former and the south and east of the latter. In habitat it is near
H. peplidifolium, but the capsule is dry, dehiscent and usually erect.
The exceptional characters (black-glandular stem, black-striped
petals, recurved fruiting pedicels) are all found only in Zimbabwe.
The leaves of the Madagascar plants tend to be relatively broader
than those on the mainland.

STUDIES IN HYPERICUM

Jacot Guillarmot ( 1 97 1 ) did not include H. wilmsii in her Flora of
Lesotho, re-identifying most of Bredell's records as//, aethiopicum;
but Killick cites them as H. wilmsii without comment.

10. Hypericum peplidifolium A. Rich., Tent.fl. abyss. 1: 9 (1847);
Oliv., Fl. trop. Afr. 1: 155 (1868); Engl., Pflanzenw. Ost.-Afr. C:
274 ( 1 895), in Mildbr., Deutsch Zentr. -Afr. Exped. 1 907- 1 908 2:
560 (1913); De Wild., PL Bequaert. 1: 242 (1922), 5: 404
(1932); Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 177
(1925); Staner in Revue Zool. Bot. afr. 23: 221 (1933), 24: 218
(1933), in Bull. Jard. hot. Etat Brux. 13: 68 (1934); Nod. in Bot.
Notiser 1934: 101 (1934); Robyns, FL Sperm. Pare Nat. Albert
1: 618 (1948); Andrews, FL Anglo-Egyptian Sudan 1: 212
(1950); Exell & Mendon9a, Consp. Fl. Angol. 1: 370 (1951);
Milne-Redh., Fl. trop. E. Afr. Hyperic.: 9 (1953); Hutch. &
Dalz., Fl. W. trop. Afr. 2nd ed. (ed. Keay) 1: 287 (1954); N.
Robson in Kew Bull. 12: 443, 445, map 2 (1958), in Exell &
Wild, Fl. Zamb. 1: 383. t. 73C (1961); Cufod. in Bull. Jard. hot.
Etat Brux. 29: Suppl.: 588 (1959); Spirlet, Contr. Fl. Congo,
Rwanda, Burundi, Guttiferae: 4 (1966); Moggi & Pisacchi in
Webbia 22: 264, f. 9, map 5 (1967); Bamps in Boutique, Fl.
Congo, Rwanda, Burundi Guttiferae: 3 (1970), in Bull. Jard.
hot. natn. Beige 41: 433 (1971), in Distr. Afr. PL 3: map 67
(1971); Agnew, Upland Kenya Wild Fls: 187 & f. (1974);
Robson in Bamps, Robson & Verdcourt, Fl. trop. E. Afr.
Guttiferae: 31 (1978); Troupin, Fl. Rwanda 1: 299, f. 63-3
(1978). Type: Ethiopia, 'Abyssinia', Quartin-Dillon & Petit s.n.
(P-lectotype, Moggi & Pisacchi, 1967).

Fig. 24C, Map 29.

H. peplidifolium var. robustum Baker f. in Trans. Linn. Soc. London,
II (Bot.) 4: 6 (1864). Types: Malawi, Mt Milangi [Mlanje],
October 1 891 (fl & fr), Whyte 143 (BMMectotype, selected here;
K!-isotype); Mt Milanji, September 1891 (fl), Whyte s.n.
(BM!-syntype).

H. peplidifolium forma ovatum Engl. in Bot Jb. 19, Beibl. 47: 40
(1894). Types: Tanzania, Kilimanjaro, Marangu, 1550 m, Volkens
697 (Bt-syntype), Volkens 698 (Bt-syntype); Usambara Mts,
Mlalo, Hoist 47 (Bt-syntype); Usambara Mts, Lutindi, Hoist
3266 (Bt-syntype).

H. peplidifolium forma pa rvifolium Engl. in Bot. Jb. 19, Beibl. 47:
40 (1894), Pflanzenw. Ost-Afr. C: 274 (1895) ['parvifolia'];
Chiov. in L.A. Savoia-Aosta, Espl. Uaba-Uebi Sceb.: 389 (1932).
Types: Tanzania, Kilimanjaro, Marangu, 2000-2500 m, Volkens
829 (Bt-syntype); Kisinika [Kifmika], 2800 m, Volkens 1157
(Bt-lectotype, selected here; BM!, K!-isolectotypes). The BM
and K specimens of this number are said to have come from
Mawenzi at 3000 m.

H. peplidifolium var. oblongifolium Engl. in Bot. Jb. 19, Beibl. 47:
40 (1894); Pflanzenw. Ost-Afr. C: 274 (1895). Type: Tanzania,
Kilimanjaro, Kibo, 3100 m, Volkens 1545 (Bt-holotype; BM!,
K!-isotypes).

H. peplidifolium var. ovatum (Engl.) Engl., Pflanzenw. Ost-Afr. C:
274 (1895) ['ovflto'], in Annali 1st. Bot. Roma 7: 20 (1898).

H. peplidifolium forma robustum (Baker f.) Engl., Pflanzenw. Ost-
Afr. C: 274 (1895) ['robusta'].

H. peplidifolium var. diestelianum Engl. in Bot. Jb. 40: 555 (1908).
Type: Cameroon, Cameroons Mtn, Buea, Diestel s.n.
(Bt-holotype).

H. peplidifolium var. ovatum forma humileRiva in sched. ['humilis']
(cf. Moggi & Pisacchi, 1967: 264).

H. peplidifolium var. anagallidifoliumChiov. in sched. (cf. Moggi &
Pisacchi, 1967: 264).

169

//. sp. C. sensu Milne Redh., Fl. trop. E. Afr. Hyperic.: 13 (1953).
H. humbertii sensu Spirlet, Contr. FL Congo, Rwanda, Burundi,

Guttiferae: 5 (1966) pro parte quoad, spec. Troupin 2669 (fide

Bamps, 1970: 3).

Icones: Moggi & Pisacchi in Webbia 22: 265, f. 9 (1967); Agnew,
Upland Kenya Wild Fls: 187 (1974).

Perennial herb, wiry or rarely woody, up to 0.6(-0.9) m tall but often
much less, 0.1-0.9 m in diam., tufted, with branches ascending to
prostrate, sometimes rooting at proximal nodes or budding from
horizontal roots, rather slender. Stems wholly terete or sometimes
slightly 2-lined above, eglandular or (in S. of range)
black-gland-dotted. Leaves with petiole 0.5-1.5 mm long; lamina
3-26 x 2-20 mm, ovate to elliptic or obovate or rarely suborbicular,
spreading; apex rounded to obtuse, margin plane, base rounded to
cuneate; venation: 2-3 pairs of main lateral veins, branching to form
rather inconspicuous tertiary reticulation; laminar glands pale and
sometimes a few black, punctiform; intramarginal glands dense,
black. Inflorescence 1 -flowered, terminal, often with 1 or 2 flower-
ing branches from uppermost 1 (2) nodes, branching rarely repeatedly
pseudo-dichotomous; pedicels 4-40 mm long, reflexed in fruit.
Flowers 8-15 mm in diam.; buds ovoid to ellipsoid, obtuse. Sepals
4-5 x 3-4 mm, imbricate, very unequal, the inner ones narrower,
obovate to elliptic or oblong or lanceolate, obtuse or apiculate to
rounded, entire; veins 3, branching and reticulating; laminar glands
pale and occasionally some black, punctiform; inframarginal glands
dense, all or mostly black. Petals bright to primrose yellow, often
tinged red, 7-8(-14) x 2-4 mm, c. 1.7 x sepals, obovate; laminar
glands pale or very rarely also black; marginal glands round most of
margin, black, sometimes also submarginal. Stamens obscurely
3(4-5 )-fascicled or irregular, 20-40(-60), longest 5-6 mm, c. 0.75 x
petals; anther gland black. Ovary 2.4-3 x 1.4-1.9 mm, ovoid to
broadly ellipsoid, acute to obtuse; styles 5(4), 2.5-4 mm long, c.

170

N.K.B. ROBSON

1-1.3 x ovary, divergent; stigmas scarcely capitate; placentae 5(4),
axile. 'Capsule' bacciform, indehiscent, 6-11 x 5-9 mm, c. 2 x
sepals, broadly ovoid to subglobose, smooth. S^dsyellowish brown,
0.6-0.8(-1) mm, not carinate; testa finely reticulate-scalaritbrm. 2n
= 16(1. &O. Hedberg, 1977).

Marshes, swamps, streamsides, roadsides, pastures, temporary leys
and abandoned cultivation in upland and moorland grassland;
1100-3750 m (W., NE and E. tropical Africa), 600-2350 m (S.
tropical Africa).

Cameroon, Fernando Poo, Ethiopia, Sudan Republic, Zaire, Rwanda,
Burundi, Uganda, Kenya, Tanzania, Malawi, Zambia, Mozambique,
Zimbabwe and Angola.

CAMEROON. Adamawa: Mambela Distr., Ngel Nyaki, 1650 m, 20
January 1958 (fl & fr), Hepper 2811 (K); Bamenda, Bambalue Forest
Reserve, 2100 m, 3 September 1952 (fl). Savory UCI 400 (K); Cameroon
Mtn, S. side, Mann's Spring, 2190m, 25 March 1947 (fl), P. W. Richards 952\
(K).

EQUATORIAL GUINEA. Fernando Poo: Moka [Mioka], 1500 m, 27
September 1959 (fl & fr), F. Melville 689 (K).

ETHIOPIA. Arssi: Galama Mts, c. 30 km ESE of Asella, c. 3 km E. of
Boraluco, Chillalo Awraja, 3750 m, 7 September 1967 (fl), Hedberg 4194
(K). Bale: Dincho. 3120m, 17 April 1970(fl), Gilbert 1781 (EA, K). Eritrea:
Hamasan, Asmara and Taclesan, 5 May 1892, Terraciano & Pappi 285 (Fl).
Gammu-Gofa: Cencia, 16 March 1938 (st), Vatova 2073 p.p. (Fl). Gojjam:
Choke Mts, vicinity of upper Ghiedeb valley, near N. peak, Arnt Makereke,
c. 3600 m, 14 August 1957 (fl), Evans & Hellier in Camb. U. Bot. Exped. 529
(K). Gondar: Simien Mts, Geech area, 3500 m, 4 September 1968 (fl), B. &
E. Nievergelt 1 166 (EA). Harerghe: c. 50 km due W. of Harar, NW face of
Gara Mullata Mtn, 2500-3000 m, 14 February 1962 (fl), Burger 1472 (K).
Illubabor: just outside Gore on Mettu road, c. 1900 m, 7 December 1970 (fl
& fr), Danish Bot. Exped. 1970 590 (C*, K). Keffa: c. 15 km NW of Bonga
on Wush-Wush road, c. 1 800 m, 1 7 August 1 965 (fl & fr), de Wilde 7775 (K,
WAG*). Shoa: Entoto Hill, just N. of Addis Ababa, 2650 m, 28 November
1972 (fl & fr),Danish Bot. Exped. 1972-73 1382 (C*, K). Sidamo: Garbicho,
nearWondo, 2400 m, 3 1 January 1954 (fl & fr), Mooney 5677 (K).Tigray: am
Rand der Aker, 1650-2400 m, 10 October 1862 (fr), Schimper 544 (BM, K,
Z).

SUDAN REPUBLIC. Equatoria: Imatong Mts, Gilo, 1800 m, 29 June
1947 (fl), MacLeay 1 17 (BM).

KENYA. Turkana: W. Suk, Kapenguria, 2100 m, 13 May 1932 (fl & fr),
Napier 1927 (K). Rift Valley: Nakuru Distr., Eastern Mau Forest Reserve,
camp 10, 2650 m, 1 September 1949 (fl), Maas Geesteranus 6016 (K, Z).
Central: Mt Kenya, N. sector, 2850 m, 30 July 1949 (fl & fr), Schelpe 2476
(BM). Nzanza: Trans-Nzoia Distr., Mt Elgon, 3360 m, 24 February 1935 (fl),
Taylor 3660 (BM). Southern: Narok Distr., Masailand, Oldevesi Lemoko, c.
72 km N. of Aitong, c. 2070 m, 28 April 1961 (fl & fr), Glover, Gwynne &
Samuel %12(K).

UGANDA. Northern: Karamoja Distr., Moroto Mtn, 2550 m, January
1959 (fl), Wilson 650 (K). Western: Toro Distr., Fort Portal, the fort, 1525 m,
12 May 1953 (fl & fr), Osmaston 2899 (K); Kigezi Distr., Kabale-Bufundi
road, 2100 m, 12 October 1929 (fl), Snowden 1502 (BM, K). Eastern:
Bugishu Distr., Sipi, 1650 m, 30 August 1932 (fl & fr), Thomas 397 (K).
Buganda: Masaka Distr., Minzilo, 30 October 1925 (fl), Maitland 1 197 (K).

ZAIRE. Lacs Edouard et Kivu: Rutshuru, mont Katale, December 1937
(fl & fr), Lebrun 9 1 85 (BR*, K, P*); Ruanoli - Lamia ridge, 2700 m, 5 August
1952 (fl), Ross 852 (BM). Lac Albert: Nioka, 1800 m, 4 July 1946 (fl), Taton
130 (BM, BR*, K); mont Korovi, 2100m, 24 June 1958 (fl & fr), Bamps2?>\
(BR*, K). Haut-Katanga: Pare National de 1'Upemba, Lusinga, de Witte 501 2
(BR*); Marungu Mts, environs de Kasiki, r. Lunangwa, 2300 m, February
1970, Lissowski, Malaisse & Symoens 10666 (EBV*).

RWANDA. Cyangugu: Kirambo, R. Karunduru, Rangiro, foret de
Nyungwe, 1600 m, 13 June 1978 (fl), Raynal 20524 (BM, P*); Ruhengeri,
Mushas, Zappelli 38 (BR*).

BURUNDI. Muramoya, Nyabiagondo, 2 1 00 m, 2 1 January 1 966 (fl &
fr), Lewalle 305 (BM, BR*, K); Ngozi, Becquet 867 (BR*).

TANZANIA. Lake: Ngara, Bushubi, Kaza, 1500 m, 15 May 1960 (fl),
Tanner 4987 (K). Northern: Kilimandjaro-Sud, c. 1800 m, 12 January 1934

(fl & fr), Schlieben453Q (BM, K, Z);Arusha Distr., Ngurdoto Crater National
Park, Leopard Point, 1620 m, 19 March 1966 (fl & fr), Greenway & Kanuri
12440 (K). Tanga: W. Usambaras, Lushoto - Shume road, Magamba forest,
1800 m, 1 March 1953 (fl & fr), Drummond & Hemsley 1362 (K). Western:
Sumbawanga Distr., Rukwa Escarpment, Nsangu, c. 2000 m, 2 January 1962
(fl), Robinson 4872 (K). Eastern: Morogoro Distr., Mgeta, Kibuko, March
1955 (fl), Semsei 2043 (K). Southern Highlands: Mbeya Distr., Mbeya Mtn,
2100m, 13December \962(f\&fr), M.Richards 17012(K);KyimbilaDistr.,
1350 m, August 1910 (fl), Stolz 263 (K, LU, Z). Southern: Songea Distr.,
Matengo Hills, Miyau, 1620 m, 2 March 1956 (fl & fr), Milne-Redhead &
Taylor 8941 (K); Ungwe-Thal, 1300 m, 25 November 1931 (fl), Schlieben
1452 (BM.Z).

MALAWI. Northern: Nyika Plateau, Lake Kaulime, 2150 m, 24 Octo-
ber 1 958 (fl & fr), Robson 338 (BM, K); Vipya, Chikangawa, 22 January 1 956
(fl), Chapman 358 (BM). Central: Dedza Mtn, 23 October 1956 (fl & fr),
Banda 291 (BM, SRGH). South: Shire Highlands, Ndurandi, December
1893 (fl & fr), Scott-Elliot 8483 (BM, K); Mt Mlanje, W. Tuchila, 1800 m, 6
July 1956 (fl & fr), Newman & Whitmore 19 (BM, K).

MOZAMBIQUE. Zambezia: Gurue, sope do pico Namuli, 24 Septem-
ber 1944 (fl), Mendonca 2257 (BM, LISJC).

ZIMBABWE. Eastern: Inyanga, Trias Hill, December 1919 (fl & fr),
Philomene i n Eyles 5181 (K, SRGH); Inyanga prope villam Inyanga Down in
convalle prope flumen Tranga, c. 1 800 m, 30 January 193 1 (fr), Norlindh &
Weimarck 4732 (BM, LU); Rhodes Inyanga Estate, 16-20 November 1931
(fl), Brain 6917 (K, SRGH).

ZAMBIA. Northern: Abercorn [Mbala], Lake Chila, 29 December 1954
(fl), M. Richards 3786 (K). Eastern: Nyika Plateau, c. 4 km from [N.
Rhodesian] Rest House, 2100 m, 22 October 1958 (fr),Robson261 (BM, K).
Western: Mwinilunga, 17 November 1937 (fl & fr), Milne-Redhead 3283
(K).

ANGOLA. Benguela: Ganda, c. 1300 m, November 1937 (fl), Pittard
99 (BM).

H. peplidifolium is variable in habit and in size of parts, but none of
the named infraspecific taxa is worthy of recognition. The tallest
plants with rather woody, ascending stems occur in the southern part
of its area (Malawi, Zimbabwe, Mozambique), and the tendency
towards 4 rather than 5 styles is greater there, too. Indeed, Mendonqa
2257 (Mozambique, Zambezia) varies towards//, natalense, whereas
Brain 6917 (Zimbabwe) has 4 styles and varies towards H. wilmsii.
In general, however, the bacciform fruit on reflexed pedicels will
distinguish H. peplidifolium from its nearest relatives.

Just as H. wilmsii has eglandular stems in South Africa and
black-gland-dotted stems in Zimbabwe, so H. peplidifolium has
black-gland-dotted stems in Mozambique and Zimbabwe and
eglandular stems elsewhere. A similar character change is found in
H. aethiopicum (sect. 27. Adenosepalum, p. 180). The significance
of this concentration of gland-dotted stems in the
Zimbabwe-Transvaal region is unclear to me.

Sect. 27. ADENOSEPALUM Spach in Annls Sci. nat.
(Bot.) II, 5:357(1836).

Shrubs, shrublets or wiry to soft herbs up to 2.5 m tall, deciduous
when woody, glabrous or with simple hairs, with dark (black) glands
nearly always on leaves (sometimes all pale in Sp. 15), usually on
sepals and anthers and sometimes on petals and stems (black or
rarely red); branching below inflorescence lateral. Stems 4-lined and
± ancipitous or 2-lined at first or wholly terete, eglandular or rarely
red- or black-gland-dotted; cortex exfoliating in strips or sheets;
bark smooth and finely striate, sometimes flaking irregularly. Leaves
opposite or very rarely and abnormally 3-whorled, decussate, sessile
or rarely very shortly petiolate, free, deciduous at base or persistent;
lamina entire or rarely undulate to serrulate with prominent glands,
with venation pinnate, open (laterals ending freely) or ± closed
(upper or all laterals incurved and uniting), with tertiary venation ±
densely reticulate; laminar glands punctiform; marginal gland-dots

STUDIES IN HYPER1CUM

171

dense or rarely ± sparse; ventral glands absent. Inflorescence \-c. 6(5) Sepals with pale and black laminar glands 22. cuisinii

200-flowered, with branching dichasial (first node) then monochasial,

,, . 0 , ... ... , ... r Sepals with only pale laminar glands 23. lanugmosum

from 1-8 nodes often with subsidiary branches from lower nodes;

bracts and bracteoles reduced, transitional in form to sepals. Flowers 7(5) Leaves (20-)30-60 mm long, the base narrowly cuneate toangustate,

stellate, homostylous. Sepals 5, free or up toe. 0.2 united, persistent, almost always with prominent marginal black glands; shrub, strag-

erect in fruit, with margin subentire to glandular-denticulate or - gling to sPreading ^low ... ... 1. glandulosum

ciliate; veins 3-7; laminar glands pale and/or black, linear to Leaves 6-20(-30) mm long, the base ± broadly cuneate to cordate

punctiform; marginal and/or inframarginal glands black, when mar- with immersed submarginal black glands; shrubs (rarely) or

ginal often flat-topped. Petals 5, persistent and twisting together subshrubs or herbs 8

round developing capsule or occasionally separately, with apiculus 8(7) Leaf apex acute to obtuse; plant a subshrub to 0 4 m with stem

present, subapical or apical, acute to apiculate, or absent, margin branched; inflorescence dense 2. abilianum

with ± prominent glands or entire; marginal glands black or absent;

laminar glands pale, linear to punctiform and/or black, punctiform Lcaf apex rounded or' if subacute to obtuse' then Plant a herb

, .-f c. s • i c •„ j T i 1 /• 4Tx branched at base, usually without stem branches; inflorescence ±
to elongate-punctiform. Stamen fascicles 5, united 2+2+1 (i.e. 3 )

or rarely 2+1 + 1 + 1 (i.e. '4') but sometimes indistinct, persistent,

totalling c. 20-70 stamens; filaments united very shortly; anthers 9<8) Petals with only marginal black glands; stem eglandular,± branched;

yellow, gland black or very rarely amber; pollen types I, X. Ovary shrubs or subshrubs or suffruticose herbs ... ... 10

with 3 loosely to completely axile placentae, each ¥-ovulate; styles Petals with laminar as well as marginal black glands; stem often

3(4), free with bases discrete; stigmas scarcely capitate or usually black-gland-dotted, usually unbranched; perennial herb, sometimes

small. Capsule 3(4)-valved, subcoriaceous to chartaceous, with with woody base (6. aethiopicum) 12

valves narrowly longitudinally vitiate. Seeds narrowly cylindric, not ,n,0> c

» fe 10(9) Sepals entire to subentire with marginal black glands, 4.5-8 mm

or scarcely connate, with apical expansion; testa almost smooth or long. jnflorescence 1-c. 25- flowered from 1-3 nodes 1 1

shallowly linear-reticulate to li near- foveol ate or finely scalariform.

Sepals black-glandular-denticulate to -ciliate, 3-3.5 mm long; inflo-
BASIC CHROMOSOME NUMBERS (X). 10, 9, 8; ploidy 2, 4. rescence c. 60-flowered from 3^ nodes 5. afrum

HABITAT. Among rocks (often calcareous) or scrub, open grass- 11(10) Leaves with base rounded to cordate-amplexicaul, sessile, (5-)7-22

land, along streams, in marshes or wet ground; sea level to 2400 m mm wide, margin undulate; slender erect shrub or subshrub with

(Europe, Mediterranean, NW Africa, Macaronesia), 1 800-3000 m branches ± strict ... ... 3. conjungens

(SW Arabia, NE Africa), 1 100-3900 m (E. Africa), 2000 m (An- Leaves with base cuneate to rounded, usually shortly petiolate, 2.5-

gola), 135-1850 m (SE Africa). 7(-8) mm wide, margin plane; ± spreading much-branched shrub

„ ,,,.,- with branches usually ± ascending 4. kiboense

DISTRIBUTION. Canary Is., Madeira; Europe N. to N. England and

S. Norway, E. to S. Finland, Poland, Byelorussia; Ukraine; Georgia 1 2(9) Stems black (rarely amber)-gland-dotted; sepals entire to subentire

and NE Turkey; SW Turkey, Cyprus and the Levant; N. Africa and with marginal glands immersed to prominent ...

adjacent Arabia, Ahoggar Mts; NE Africa and adjacent Arabia 6a. aethiopicum subsp. sonderi

(Asir); E. and SE Africa; Angola. Stems eglandular; sepals with margin distally or wholly black-

elandular-denticulate to -ciliate ....

24 species (+ 3 subspecies and 2 hybrids). ... . ... .

6b. aethiopicum subsp. aethiopicum

Key to sects 27. Adenosepalum and 28. Elodes

1 Plant completely glabrous 2

Plant with indumentum on at least young stem or leaves 13

2(1) Bracts and bracteoles densely glandular-auriculate 3

Bracts and bracteoles without gland-fringed auricles 4

3(2) Inflorescence curved-pyramidal or corymbiform to capitate, all or at
least partial inflorescences dense; stem and lamina of leaves and
sepals without black glands 17. montanum

Inflorescence broadly pyramidal to subcorymbirbrm, lax; stem and

lamina of leaves and/or sepals with black glands

18b. annulatum subsp. intermedium

4(2) Plant a shrub with leaves triangular-lanceolate, acute to subacute, ±

deeply cordate-amplexicaul

16. reflexum'

5(4)

Plant a shrub with leaves not having this combination of characters
or a subshrub or herb 5

Plant a low herb with stems weak, decumbent or diffuse; sepals
obtuse to rounded 6

Plant a shrub or subshrub or herb with stems wiry, erect to decum-
bent; sepals aristate or acute to rounded (subsect. \.Aethiopica)

... 1

'See also lead 15.

13( 1 ) Bracts and bracteoles entire to regularly glandular-ciliate but with-
out auricles, or if glandular-auriculate then leaf pairs connate .. 14

Bracts and bracteoles densely glandular-auriculate or with crowded
longer-stalked glands at base; leaf pairs free (subsect. 4.
Adenosepalum excl. Sp. 16) 25

14(13) Plant a shrub; stem tomentose to puberulous; leaves glabrous or
rarely villous-pilose 15

Plant a perennial herb; stem and leaves both with indumentum
16

15(14) Leaves glabrous, entire, base cordate to rounded; stem densely
villous-tomentose 16. reflexum

Leaves villous-pilose and/or with protruding marginal glands and/or
base cuneate; stem glabrous or ± densely villous-tomentose
Ix. x joerstadii

16(14) Flowers pseudo-tubular; stamen filaments in each fascicle c. 0.65
united; sepal marginal glands reddish (sect. 28. Elodes)
1. elodes

Flowers stellate; stamen filaments in each fascicle basally united;
sepal marginal glands black 17

17(16) Leaf pairs free; inflorescence usually lax (subsect. Pubescentes)
18

Leaf pairs all or mostly connate; inflorescence ± congested (subsect.
Caprifolia) 23

172

N.K.B. ROBSON

18(17) Sepals (dorsally) and inflorescence branches villous or tomentose
19

Sepals and upper parts of inflorescence (at least above bracteoles)
glabrous 20

19(18) Sepals 5-10 mm long, linear-lanceolate to lanceolate, apex long-
(usually eglandular-) aristate, margin entire to subentire
7. pubescens

Sepals 3-5(-6) mm long, lanceolate to ovate or broadly elliptic,
apex acute to shortly aristate and usually glandular, margin with
prominent sessile glands or glandular cilia 9. tomentosum10

20(18) Stem spreading to appressed-puberulous; leaves papilliform-
puberulous 8. psilophytum

Stem pubescent to tomentose with spreading hairs; leaves pubescent
to tomentose 21

21(20) Sepals and petals with only pale laminar glands ... 10. somaliense
Sepals and petals usually with some black laminar glands 22

22(2 1 ) Stems erect, not rooting; leaves triangular-lanceolate to narrowly
oblong, at least lower condensed-tetrastichous; flowers c. 25 mm in
diam. ... ... 11. collenettiae

stem glandular or not; stem and leaves glabrous to densely pubes-
cent 30

30(29) Petals red-tinged dorsally; stem usually densely black (rarely red) -
gland-dotted; leaves and sepals sometimes with laminar black glands
18c. annulatum subsp. afromontanum

Petals not red-tinged (rarely red-veined) dorsally; stem eglandular
or rarely sparsely black-gland-dotted; leaves and usually sepals

without laminar black glands

18b. annulatum subsp. intermedium

3 1 (27) Stems erect to ascending, strigose-pubescent; leaves sessile, 1 2-35
mm long 19. delphicum

Stems procumbent, weak, pilose; leaves petiolate, 8-15 mm long
20. athoum

32(26) Sepals with laminar black glands, margin always glandular-ciliate
33

Sepals without laminar black glands, margin sometimes eglandular
and denticulate or entire 23. lanuginosum

33(32) Sepals 3.5-5 mm long, margin long-glandular-ciliate (stalk more
than 2 x gland); leaves sessile, 15-55 mm long; stems erect to
decumbent but not rooting 21. atomarium

Stems ascending to prostrate, sometimes rooting; leaves narrowly
oblong to elliptic or oblanceolate, neither condensed nor manifestly
tetrastichous; flowers 9-13 mm in diam 12. sinaicum

Sepals 2.5-3.5 mm long, margin medium- to short-glandular-ciliate
(stalk 2 x gland or less); leaves sometimes petiolate, 2-15(-23) mm

long; stems decumbent to diffuse-ascending, rooting

23(17) Sepals acute to subacuminate, mostly oblong to elliptic, marginal 22. cuisinii

glands sessile or some on very short cilia 24

Subsect. 1. Aethiopica N. Robson in Bull. nat. Hist. Mm.

Sepa s aristate, narrowly lanceolate, with marginal elands all on T j m *.\ *** *t\ /inr>->\ <-r IT i • • TT- L
- { . .e ,. Lond. (Bot.) 23: 69 (1993). Type: H. aethiopicum Thunb.

short to long cilia or fimbnae 15. caprifohum J*

(see p. 90).

24(23) Leaves broadly elliptic to suborbicular, finely bullate, lower pairs

(always?) free 13. coadunatum Shrubs or subshrubs or perennial herbs, completely glabrous; leaves

free; bracts and bracteoles not glandular-auriculate. Species 1-6.
Leaves ± broadly elliptic or broadly oblong to obovate, not bullate,

all pairs connate 14. naudinianum

1. Hypericum glandulosum Aiton, Hort. kew. 3: 107 (1789), 2nd
25(13) Leaves scabrid beneath; plant otherwise glabrous ed 4. 428 (1812);Vahl, Symb. Bot. 2: 86 (1791); Willd., Sp. pi. 3:

1464 (1802); Buch in K. Akad. Wiss. Berlin Abh. 1816-1817:

Leaves pruinose or scabrellous to pubescent or villous on both sides; 366, 380 (1817); Choisy, Prodr. monogr. Hyperic.: 53 (1821), in

stem pruinose or scabrellous to pubescent or pilose or rarely (22. DC., Prodr. 1: 551 (1824); Link in Buch, Phys. Beschr. Canar.

cuisinii and 23. lanuginosum both in part) glabrous 26 Ins : 1 53 j 66 ( 1 828); Webb & Berth., Phytogr. canar. 1: 44, t. 3

26(25) Bracts and bracteoles densely glandular-auriculate 27 ( 1 836); Spach in Annls Sci. nat. (Bot.) II, 5: 357 ( 1 836); Lowe,

Fl. Madeira: 76 ( 1 868); Masferrer in An. Soc. esp. Hist. nat. 9: 27
Bracts and bracteoles not auriculate but often with longer glandular j p. ^ & p^^ ^ ^ Canaries, m(} 909); Menezes,

cilia towards base 32 . ~n,,n,A^ c. t or- /- i A i c- ;

Fl. Archip. Madeira: 29 (1914); Stefanoff in God. Agr.-les. Fak.

27(26) Stem and leaves puberulous to pubescent; rootstock not creeping or Univ. Sofiya 11: 148 (1933), 12: 82 (1934), in Pflanzenareale

rooting; stem sometimes gland-dotted; leaves, sepals and petals 4( ] ): Karte 2b ( 1 933); Grabham, Pis seen in Madeira: 95 ( 1 934);

sometimes with laminar black glands 28 Ceballos & Ortuno, Veg. Fl.for. Canar. occid.: 387 (1951); Lid in

Stem and leaves strigose-pubescent to pilose or villous; rootstock Skr. norske Vidensk.-Akad. I, Math.-Nat. Kl., N.S. no. 23: 120

creeping and rooting; stem, leaves, sepals and petals without laminar (1967); Bramwell, D. & Z., Wildfls Canary Is.: 162, f. 42, t. 199

black glands 31 (1974); Voggenr. in Dissert. Bot. 26: 655, 689 (1974); Kunkel,

28(27) Sepals acute to subacute, (4-)5-8 mm long; petals (8-)10-15 mm Endemismos Canarios: 294 (1977); N. Robson in Cullen et al.,

long; stems 0.2-0.75 m long, if decumbent then not rooting (18. Eur. Gdn FL 4: 72 (1995>- TyPe: Madeira, without precise

annulatum) 29 locality, Cook's First Voyage, fl. 1768, Banks & Solander s.n.

(BM !-holotype). Dryander (the actual author ofHortus kewensis)

Sepals rounded, 2.6-4.5 mm long; petals 6-8 mm long; stems 0.04- dtes om <Introduced , 777 Fr Masson: . but there is no Masson

0.15 m long, it decumbent then rooting 24. decaisneanum . . . , .

specimen in the BM collection and Masson s introduction may

29(28) Petals, sepals and leaves without laminar black glands; stem have been of living material only. At any rate, Dryander's de-

eglandular; stem and leaves densely pubescent scription was almost certainly derived from the Banks & Solander

... 18a. annulatum subsp. annulatum specimen, which therefore should be regarded as the holotype.

Petals and sometimes sepals and leaves with laminar black glands; Fig. 25 A, Map 30.

10 Plants with characters intermediate between Spp. 7 and 9 are probably hybrid (see
p. 184).

Icon: Webb & Berth., Phytogr. canar. 1: t. 3 (1836).

Shrub 0.25-c. 1 .4 m tall, straggling to spreading below, with branches

STUDIES IN HYPERICUM

173

18

16

Map 30 Sect. 27: 1. H. glandulosum • specimens, O record. Limits on Tenerife and La Palma according to Voggenreiter (1974).

rigid above, the whole plant glabrous. Stems pale yellowish to
reddish, shallowly 4-lined but not ancipitous when young, soon
terete; internodes shorter than leaves; cortex exfoliating in strips;
bark straw-brown, finely striate. Leaves sessile; lamina (20-)30-60
x (6-) 10-20 mm, ± narrowly elliptic to elliptic-oblanceolate, some-
what paler beneath, chartaceous, not glaucous, plane, spreading;
apex acute or subapiculate to obtuse or rounded, margin entire or
undulate to serrulate, with prominent but sessile round- or flat-
topped black glands or proximally or rarely wholly entire with
glands immersed, base narrowly cuneate to attenuate; venation: (2)3
pairs of laterals curved-ascending from lower 0.35-0.4 of midrib,
tertiary reticulation obscure; laminar glands pale, dense, unequal, ±
prominent; marginal and/or intramarginal glands black, dense.
Inflorescence c. 10^5-flowered from up to 3 nodes, curved-
corymbiform to hemispherical, dense; pedicels 3^.5 mm; bracteoles
(and bracts at uppermost 2 nodes) reduced, narrowly elliptic, with

prominent marginal black glands. Flowers c. 15-20 mm in diam.;
buds narrowly ellipsoid, acute. Sepals 4.5-7 x 1.5-2(-2.5) mm,
slightly unequal, free, elliptic-oblong to elliptic or (the smaller)
lanceolate-elliptic, acute, with sessile marginal glands or distally
glandular-denticulate; veins 5, branching; laminar glands pale or a
few black, linear or interrupted; marginal glands black, ± flat-
topped, prominent, dense. Petals rather pale dull yellow, tinged or
lined bright red dorsally, 10-14 x 3^4 mm, c. 2 x sepals, narrowly
elliptic-oblong, acute (i.e. apiculus acute, apical); laminar glands
pale, striiform to punctiform; marginal glands all black or some pale,
all prominent or some immersed distally. Stamens c. 25-30, longest
7-9 mm, c. 0.7 x petals; anther gland amber. Ovary 2.5-3 x 1 .5 mm,
ellipsoid; styles 5-6 mm, 2 x ovary, spreading-incurved. Capsule 4-
5.5 x 2.5^4- mm, ovoid, shorter than sepals, enclosed by petals
twisting together. Seeds pale yellow-brown, 0.5-0.6 mm long; testa
very shallowly linear-foveolate, almost smooth. 2n=18 (van Loon &
deJong, 1978;Reynaud, 1986), 40 (Dalgaard, 1986).

Open rocky hillsides or cliffs in Laurus forest or among Erica
thickets; 300-900 m or sometimes down to maritime region in
barrancos (Canary Is.), 200-700 m (Madeira).

Canary Islands (Tenerife, Gomera, La Palma, Gran Canaria), Ma-
deira.

CANARY ISLANDS. Tenerife: Vueltas de Taganana, 1 6 July 1 858 (st),
Lowe Ten 183 (BM, K), ibid., 12 April 1971 (fl), Bramwell & Humphries
3395 (BM); Mercedes, 5 June 1892 (fl), Murray s.n. (BM, K), loc. cit., 20
March 1936(fl), Nielsen 1697 (C*, K);Anagagebirge zwischen El Bailladero
und Roque Chinobre, 700 m, 12 January 1977 (fl), Gilli s.n. (W); Montanas
de Anaga, Barranco de las Huertas, 13 km above San Andres, 8 April 1975
(fl),/, M. & P. Cannon 4677 (BM). Gomera: El Monte, Hermigua, 18 April

174

N.K.B. ROBSON

Fig. 25 A. H. glandulosum: (a) habit; (b) stem node; (c) leaf; (d) flower; (e) sepal; (f) petal; (g) closed flower, showing twisted petals; (h) capsule. B. H.
conjungens: (i) inflorescence; (j) section of stem, showing flaking bark. C. H. kiboense: (k) leaf. D. H. abilianum: (1) leaf. E. H. afrum: (m) leaf (a, i x '/i;
c x 1; d, k-m x 2; b, d, e-h, j x 3). A. Jarvis 628. B. Robson 269. C. Rogers 155. D. Exell & Mendortfa 2995 A. E. Robert & Cosson in Herb. Soc. Dauph.
5205.

STUDIES IN HYPERICUM

175

1861 (fl), Lowe 146 (BM, K); near Vallehermose, Chorros de Epina, 700 m,
9 May 1977 (fl), Jarvis 628 (BM). La Palma: Los Sauces, Barranco de Agua,
25 May 1858 (fl), Lowe P.2 10 (BM); Barranco Galga, 23 February 1888(fl),
Kuntze s.n. (K). Gran Canada: no specimens seen.

MADEIRA. Curral das Freiras, 3 July 1 855 (fl), Lowe s.n. (BM, K), 1 2
July 1974 (fl & fr), McClintock s.n. (BM); near Machico, 300 m, 28 April
1924 (fl), Riley 26 (BM, K), Levada Machico, between valleys of Ribeiras
Funda and Seca, c. 200 m, 4 April 1985 (fl), Press 710 (BM).

H. glandulosum has as its nearest relative in sect. 1 . Campylosporus
the east African H. quartinianum, its flower and fruit differing
essentially from those of the latter in having '3' stamen fascicles, a
3-merous ovary with spreading styles, fading petals that twist around
the developing ovary, and almost smooth seeds. The inflorescence
branching is initially dichasial, as it often is in the closely related H.
roeperianum, which has a more widespread African distribution
than H. quartinianum (see Robson, 1985: 194-201).

H. glandulosum is rather distantly related to the other Canary
Island shrub in sect. Adenosepalum, 16. H. reflexum, and differs in
its glandular-margined leaves (which have laxer venation and are
broadest about the middle, not below), and glandular-ciliate sepals
and bracteoles. In addition, H. glandulosum includes tetraploids on
the base 10 (2n=40) and diploids on the base 9 (2n=l 8), whereas H.
reflexum is apparently diploid only (2n=18). H. glandulosum is
usually completely glabrous, whereas H. reflexum normally has a
pubescent stem. Plants intermediate in leaf form between the elliptic
typical of//, glandulosum and the triangular-lanceolate of typical//.
reflexum have been found. Those like H. reflexum but with ovoid or
ovoid-lanceolate leaves have been named H. reflexum var.
myrtillifolium Bornm., whereas those similar to H. glandulosum in
leaf shape but with villous-tomentose stems and villous-pilose
leaves have been described as a distinct species, H. joerstadii Lid.

It should be noted that: i) H. glandulosum and H. reflexum are
typically not only distinct but very different in stem, leaf and sepal
characters; ii) //. glandulosum in Madeira (in the absence of H.
reflexum) shows no tendency to vary towards that species; and iii)
each species has apparently given rise to quite distinct species
groups on the African mainland. For these reasons it seems best to
regard H. glandulosum and H. reflexum as 'good' species and all or
most of the variation observed in the Canaries as due to hybridiza-
tion between them.

Ix. Hypericum x joerstadii Lid in Skr. Norske Vidensk.-Akad. I.
Math.-Nat. Kl., N.S. no. 23: 121, f. 9a (1967). Type: Canary
Islands, Tenerife, Pedro Alvarez, 1954 (fl), L/ds.n. (O-holotype).

Icon: Lid in Skr. Norske Vidensk.-Akad. I. Math.-Nat. Kl., N.S. no.
23: 121, f.9a (1967).

Shrub intermediate in characters between 1 . H. glandulosum and 1 6.
H. reflexum. Stem terete when mature, glabrous or ± densely villous-
tomentose, especially when young. Leaves 30-50 x 15-30 mm,
elliptic to elliptic-oblong or obovate, glabrous or villous-pilose on
both sides, acute to rounded, margin entire or with sessile black
glands, base cuneate to rounded, tertiary reticulate venation obscure
to marked and rather dense. Sepals with black marginal glands all
prominent or some immersed and interspersed with glandular cilia.

Canary Islands (Tenerife, La Palma).

CANARY ISLANDS. Tenerife: Cumbre above Afur, 1 957, Joerstad s.n.
(O*); El Rosario S. of Esperanza, 1020 m, 1957, Lid s.n. (O*); Valle de
Guerra to Cruz Chiquita, 1 April 1975 (fl), J., M. & P. Cannon 4429 (BM);
ibid., J., M. & P. Cannon 4450 (BM); 1.6 km beyond Taganana towards
Puerto Anaga, 15 July 1858 (fr), Lowe Ten 280 (BM, K). La Palma: Barranco
del Carmen, 1 June \9\3(fl),Sprague &Hutchinson 174(K);BarrancoAgua

in Los Sauces, 220 m, 1954, Lid s.n. (O*); Barranco Santa Lucia, 200 m,
1954, Lid s.n. (O*).

Of the specimens cited above, Cannon 4429 and 4450 are typical //.
glandulosum except for the hairs on young shoots, Lowe Ten. 280 is
typical H. reflexum (and apparently with good seed) except for the
elliptic-oblong leaves, while the remainder are intermediate (except
for the hairy leaves) and are all paratypes of H. x joerstadii. It is
possible that 16. H. reflexum var. leiocladum (i.e. H. reflexum with
glabrous stems) should be included here, in which case the first lead
of couplet 4 in the key (p. 171) would indicate H. x joerstadii instead
of H. reflexum.

2. Hypericum abilianum N. Robson in Bolm. Soc. broteriana II,
53: 114, ff. 1, 2 (1980). Type: Angola, Huila, near Posto
Zootecnico, near Humpata, c. 2000 m, 21 June 1937 (fi), Exell &
Mendonca 2995 A (BM!-holotype; COI, LISJC).

Fig. 25D, Map 31.

H. kiboense sensu N. Robson in Kew Bull. 12: 437 ( 1 958), pro parte
quoad specim. Exell & Mendonca 2995A.

Icon: Bolm Soc. broteriana II, 53: 121, f. 2 (1980).

Subshrub 0.4 m tall, erect, with branches strict, glabrous. Stems
orange to vinous red, 4-lined and ancipitous when young, soon 2-
lined, eventually terete, internodes shorter than leaves; cortex
exfoliating in strips; bark reddish brown, finely striate. Leaves
sessile or with petioles to 0.2 mm long; lamina 12-16 x 3.5-6 mm,
lanceolate to narrowly oblong or elliptic-oblong, paler beneath,
chartaceous, not glaucous, with margins plane or recurved (in
drying?), spreading; apex acute to apiculate or obtuse, margin entire,
base cuneate to subcordate; venation: 3(4) pairs of laterals curved-
ascending from lower 0.3-0.35 of midrib, tertiary reticulation dense;
laminar glands pale, dense, unequal, not prominent; intramarginal
glands black, dense. Inflorescence c. 3-5-flowered from up to 3
nodes, congested, curved-corymbiform, with irregularly disposed
flowering branches from lower nodes; pedicels 1-3 mm; bracteoles
reduced, narrowly lanceolate, with dense marginal black glands.
Flowers c. 15 mm in diam.; buds ellipsoid, subacute. Sepals 6-7 x
1.4-1.6 mm, subequal, free, narrowly oblong to lanceolate, acute,
entire; veins 5-7, branching; laminar glands pale, linear, distally
interrupted to striiform; inframarginal glands black, dense. Petals
golden yellow, tinged red dorsally, 8-10 x 4-4.5 mm, c. 1 .7 x sepals,
oblong-oblanceolate, rounded with apiculus apiculate, subapical;
laminar glands mostly pale, linear to striiform or punctiform, occa-
sionally black, solitary, punctiform; marginal glands black, slightly
prominent. Stamens c. 70, longest 7-8 mm, c. 0.85 x petals; anther
gland black. Ovary c. 3 x 1.5 mm, ovoid-pyramidal; styles 4.5 mm,
1 .5 x ovary, spreading-incurved. Capsule and seeds unknown.

'Among grass and low ericoid shrubs'; c. 2000 m.

Angola (Huila).

ANGOLA. Huila: near Humpata, near Porto Zootecnico, 21 June 1937
(fl), Exell & Mendonfa 2995A (BM, COI*, LISJC*).

Map 31 Sect. 27: 2. H. abilianum *; 3. H. conjungens D; 4. H. kiboense •; 5. //. o/rwrn all records A.

STUDIES IN HYPERICUM

H. abilianum is closely related to the East African 3. H. kiboense and
4. H. conjungens, but differs from both in its lanceolate to oblong,
acute to obtuse leaves. It is nearer H. conjungens in its erect, strict-
branching habit, and its leaves are intermediate in size between those
of H. conjungens and H. kiboense but nearer in shape to those of H.
glandulosum.

H. abilianum is known so far from only one collection.

3. Hypericum conjungens N. Robson in Kew Bull. 13: 397 ( 1 959),
in Exell & Wild, Fl. zamb. 1: 381 (1961); Spirlet, Contr. Fl.
Congo, Rwanda, Burundi, Guttiferae: 6 (1966); Bamps in Bou-
tique, Fl. Congo, Rwanda, Burundi, Guttiferae: 5 (1970), in Bull.
Jard. hot. natn. Beige 41: 436 (1971), in Distrib. Plantarum
Africanarum 3: map 69 (1971); Lisowski, Malaisse & Symoens
in Bolm. Soc. broteriana II, 44: 231(1 970); Bamps, Robson &
Verde, in Polhill, Fl. trap. E. Afr., Guttiferae: 30 (1978); Robson
in Bolm. Soc. broteriana II, 53: 1 15, f. 1 (1980) ['conjugens'].
Type as for H. conjunctumN. Robson (1958) non Y. Kimura
(1938).

Fig. 25B, Map 31.

H. sp. B sensu Milne-Redh. in Turrill & Milne-Redh., Fl. trap. E.
Afr., Hypericaceae: 12(1953).

H. conjunctum N. Robson in Kew Bull. 12: 437, map 1 (1958), non
Y. Kimura (1938). Type: Tanzania, S. Highlands Prov., Njombe
Distr., Mdapo, March 1954 (fl), Semsei 1643 (K!-holotype; PRE).

H. milne-redheadii Gilli in Annln naturh. Mus. Wien 74: 425, t. 1 f.
1 (1970). Type: Tanzania, S. Highlands Prov., Njombe Distr., bei
Madunda in Livingstonegebirge, 2050 m, 29 July 1958 (fl & fr),
Gilli 176(W!-holotype).

Icon: Gilli in Annln naturh. Mus. Wien 74: 425, t. 1 f. 1 (1970).

Shrub or subshrub c. 0.3-2 m tall, slender, erect, with branches ±
strict, glabrous. Stems orange to vinous red, 4-lined and ancipitous
when young, soon 2-lined, eventually terete, internodes mostly
equalling or exceeding leaves; cortex exfoliating in sheets; bark
smooth, finely striate. Leaves sessile or sometimes lower ones very
shortly petiolate; lamina 10-30 x (5-)7-20(-22) mm, oblong to
elliptic-oblong or rarely obovate, paler beneath, chartaceous, not
glaucous, spreading; apex rounded to retuse, margin ± undulate,
base rounded to cordate-amplexicaul; venation: (2)3 pairs of laterals
curved-ascending from lower 0.2-0.25 of midrib, tertiary reticula-
tion well-developed, dense; laminar glands pale, dense, unequal, not
prominent; intramarginal glands black, dense. Inflorescence c. 5-
20-flowered, from 1-2 nodes, congested, curved-corymbiform, with
branches from up to 3 nodes below; pedicels 1-3 mm; bracteoles
foliar or usually reduced, narrowly oblong to narrowly triangular-
lanceolate, with dense marginal to inframarginal black glands.
Flowers c. 15-20 mm in diam.; buds ellipsoid, subacute. Sepals 5-
8x1-2 mm, subequal, free, oblong-lanceolate to lanceolate, acute,

177

entire to subentire; veins 5, sometimes branching; laminar glands
pale, linear, distally interrupted; marginal to inframarginal glands
black, dense. Petals primrose yellow, tinged and sometimes veined
red dorsally, 10-12 x 2-2.5 mm, 1.5-2 x sepals, oblong, rounded,
with apiculus obsolete, subapical; laminar glands pale, linear to
dorsally striiform or subpunctiform, occasionally black, subapical,
punctiform; marginal glands black, not or slightly prominent. Sta-
mens c. 80, longest 7-8 mm, 0.7-0.8 x petals; anther gland black.
Ovary 2.5-3 x 1.5-2 mm, ovoid-pyramidal; styles 2.5-3.5 mm,
(1-)1.2 x ovary, spreading-incurved. Capsule 6-7 x 3-3.5 mm,
ovoid-pyramidal, shorter than sepals, surrounded by erect, individu-
ally twisted petals. Seeds dark brown, 0.7-1 mm long; testa
scalariform-reticulate.

Upland grasslands, grassy valleys and forest margins; 1800-2910
m.

Zaire (Shaba), Tanzania (Southern Highlands: Mbeya and Njombe
districts), Malawi (North: Nyika), Zambia (Eastern: Nyika).

ZAIRE. Shaba: Marungu, Kisinde, Dubois 1090 (BR*); Marungu,
Kasiki, 2200 m, June 1969, Lisowski, Malaisse & Symoens 6476 (BR*).

TANZANIA. Southern Highlands: Mbeya Distr, Poroto Mts, Upper
Kiwira R., 1950 m. May 1938 (fl), Maclunes 415 (BM); Mbeya distr., 19.2
km from Mbeya to Chunga, 2400 m, May 1959 (fl), Procter 1215 (K);
Njombe Distr., Matamba, 1800 m, 8 January 1957 (fl), Richards 7593 (K).

MALAWI. North: Rumphi Distr., Nyika Plateau, near Nganda road, 8
June 1960 (fl), Chapman 748 (BM, K, SRGH*); Chitipa Distr., Nyika
Plateau, north end, 4 km from Nganda, 2380 m, 29 July 1972 (fl & fr),
Brummitt & Synge WC 75 (K).

ZAMBIA. Eastern: Nyika Plateau, c. 4 km SW from [Northern Rhode-
sian] Rest House, 2100 m, 22 October 1958 (fl), Robson 269 (BM, K, LISC,
SRGH).

H. conjungens is intermediate not only between 4. H. kiboense and
6. H. aethiopicum (the origin of the specific epithet), but also
between 3. H. abilianum and H. kiboense. Like the other species in
this group (Spp. 1-6), it is thus clearly a relict of considerable
antiquity; although the morphological differences between Spp. 2, 3
and 4 are not great, the geographical distances between them are. H.
conjungens differs from H. abilianum mainly in leaf shape and
internode length, whilst the shrubbier habit and more congested
inflorescence (inter alia) distinguish it from H. aethiopicum. It is
usually distinguished from H. kiboense by its longer, sessile leaves
and slender, strict habit; but the population of that species from Mt
Meru (N. Tanzania) and the specimen (Rammell F.D. 3492 (K)) from
Narok (S. Kenya) originally included by me in//, conjungens (H. sp.
A sensu Milne-Redhead, 1953), narrow the morphological gap
between these species considerably. In Fl. Trop. E.Africa, Guttiferae
(Bamps, Robson &Verdcourt, 1978: 30), I expressed doubts as to the
exact identification of 'sp. A'; but it is now clear to me that it exhibits
an extension of variation of//, kiboense (q.v.) rather than constitut-
ing a disjunct population of H. conjungens.

4. Hypericum kiboense Oliv. in Trans. Linn. Soc. II, Bot. 2: 329
(1887); Engl. in Phys. Abhand. Konigl.Akad. Wiss. Berlin 1891:
307 ( 1 892), Pflanzenw. Ostafr. C: 274 ( 1 895), Veg. derErde 3(2):
500 ( 1 92 1 ), T.C.E. Fr. in Notizbl. hot. Cart. Berlin 8: 566 ( 1 923);
Brenan, Tang. Terr. Checklist 2: 249 (1949); Milne-Redh. in
Turrill & Milne-Redh., Fl. trop. E. Afr. Hyperic.: 6 (1953);
Hedberg in Symb. Bot. Upsal. 15(1): 131 (1957); N. Robson in
Kew Bull. 12: 437 (1958); Dale & Greenway, Kenya trees &
shrubs: 235 (1961); Agnew, Upland Kenya wild/Is: 186 (1974);
Bamps, Robson & Verde, in Polhill, Fl. trop. E.Afr. Guttiferae: 30
(1978). Type: Tanzania, Kilimanjaro, 3900 m, 1884 (fl), H.H.
Johnston 136 (K!-holotype).

178

Fig. 25C, Map 31.

H. kiloense sensu H.H. Johnston, Kiliminjam Exped. App.: 338

(1886), sphalm.
H. sp. A sensu Milne-Redh. inTurrill & Milne-Redh., Fl. trap. E.Afr.

Hyperic: 12(1953).
H. conjunctum sensu Agnew, Upland Kenya wild/Is: 187 (1974).

Shrub orsubshrub up to 2.5 m tall, spreading to straggling and often
much branched, with branches ± strict to ascending, glabrous. Stems
orange to vinous red, 2-lined and ancipitous when young, soon
terete, internodes mostly shorter than leaves; cortex exfoliating in
sheets or strips; bark irregular, flaking. Leaves up to 0.7 mm
petiolate or rarely sessile; lamina 6-18(-21) x 2.5-7(-8) mm,
oblong to elliptic-oblong or obovate, paler beneath, chartaceous, not
glaucous, plane, spreading; apex rounded or mucronulate, margin
plane or rarely undulate, base cuneate to rounded; venation: (2)3
pairs of laterals curved-ascending from lower c. 0.35 of midrib;
tertiary reticulation well developed but rather obscure; laminar
glands pale, dense, unequal, sometimes prominent, sometimes also
1-2 black, subapical, punctiform; intramarginal glands black, ±
dense. Inflorescence \-c. 10-flowered, from up to 3 nodes, not or
scarcely congested, broadly pyramidal when fully developed, with
branches from up to 2 nodes below; pedicels 2-4 mm; bracteoles
reduced, lanceolate, with ± dense inframarginal black glands. F low-
ers c. 15-22 mm in diam.; buds ellipsoid, subacute. Sepals
(occasionally 4), 4.5-6 x 1-2.2 mm, subequal to equal, free, oblong-
lanceolate to lanceolate, acute, entire or usually with prominent
marginal glands or partly glandular (rarely eglandular) -denticulate;
veins 5, not branching; laminar glands pale, linear, distally inter-
rupted to punctiform; marginal to inframarginal and occasionally
submarginal glands black or occasionally reddish, dense. Petals
(occasionally 4) golden yellow, tinged red dorsally, 8-11 x 3^4.5
mm, c. 2 x sepals, elliptic-oblong to oblanceolate, rounded, with
apiculate obsolete, subapical; laminar glands pale, linear to distally
subpunctiform; marginal and sometimes inframarginal glands black,
subprominent on inner margin. Stamens c. 40-50, longest 6-10 mm,
c. 0.8-0.9 x petals; anther gland black. Ovary 3(4)-locular, 3-4 x 1-
2 mm, ovoid-pyramidal to ovoid; styles 3(4), 3.5-5, 1-1.3 x ovary,
spreading-subincurved. Capsule 6-7 x 3.5 mm, ovoid-pyramidal to
ovoid, exceeding sepals, not enclosed by spreading, individually
twisted petals. Seeds yellow-brown, c. 0.7 mm long; testa finely
scalariform-foveolate.

Upland grassland, bushland and forest margins, often along streams;
( 1800)21 00-3900 m.

Uganda (Mt Elgon), Kenya (Mt Elgon, Aberdares, Mt Kenya, Mt
Nyiru), Tanzania (Kiliminjaro, Mt Meru, Mt Oldeani, Lemagrut).

UGANDA. Eastern: Mbale Distr., Mt Elgon above Sipf, 2700 m, Sep-
tember 1934 (fl), Synge 1065 (BM); Mt Elgon, Bulambuli, 2700 m, August
1929 (fl), Sounders & Hancock 60 (K).

KENYA. Northern Frontier Distr.: Mt Nyiru, 2700 m, 5 January 1959
(fl), Newbould 345 1 (K). Rift Valley: Trans-Nzoia Distr., Mt Elgon, 3360 m,
23 February 1934 (fl & fr), Taylor 3614 (BM); Elgeyo Distr., Cherangani
Hills, Embobut Forest above Arror valley, 3240 m, January 197 1 (fl), Tweedie

N.K.B. ROBSON

3916 (K); West Aberdares, 2400 m, 29 July 1913 (fl), Battiscomhe 7 10 (K).
Central: Fort Distr., S. Kinangop Forest Reserve, Wamunu, 2400 m, 23
November 1959 (fl), Kerfoot 1415 (K); North Nyeri Distr., Mt Kenya,
Sirimon Track just N. of Timau, 2850-3000 m, 23 December 1963 (fl),
Verdcourt 3459 (K). Masai: Masai Distr., N. of Narok, c. 1800 m, n.d. (fl &
fr), Rammell F.D. 3492 (K).

TANZANIA. Northern: Moshi Distr, Kiliminsharo, S.O. Seite, 2300 m,
28 December 1933 (fl), Schlieben 44 1 9 (BM, K, P*, Z); Loliondo, Ngosaro
Samba, near Kenya border, 6 July 1956 (fl), Williams 701 (K); Arusha distr.,
Mt Meru, NE rim of crater, 2530 m, 6 October 1977 (fl), Raynal 19442 (K);
Masai distr., Lemagrut Mtn, S. side, near road to Endulea, c. 2550 m, 19 June
1965 (fl), Herlocker 129 (K); Mbulu distr., Mt Oldeani W. side, 2850 m, 16
February 1961 (fl), Newbould 5709 (K).

H. kiboense is a higher-altitude derivative (or relative) of H.
conjungens, differing essentially from it in the shrubbier straggling
habit, smaller ± petiolate plane-margined leaves, laxer inflores-
cence, more prominently glandular-denticulate sepals and fewer
stamens. The more southern populations in Tanzania (on Mts Meru,
Oldeani and Lemagrut) are also morphologically nearer to H.
conjungens; but the nearest morphologically is Rammell F.D. 3492,
from Kenya, Masai Distr. (Sp. A in Milne-Redhead (1953)). This
grew at a lower altitude than the other populations and has somewhat
larger leaves, rounded to subcordate at the base, and more numerous
stamens. It cannot, however, be included in H. conjungens because
the leaves are too narrow and mostly shortly petiolate, and is best
treated as an aberrant population of H. kiboense.

5. Hypericum afrum Lam., Encycl. 4: 166 (1797); Desf., Fl.
atlant. 2:215(1 798); Choisy, Prodr. monogr. Hyperic. : 50 ( 1 82 1 ),
in DC., Prodr. 1: 549 (1824); Batt. &Trab., Fl.Algerie, Dicots 1:
181 (1888); Barratte in Coss., Ill.fl. atlant. 2: 1 1, t. 102 (1892);
Julien, Fl. Constantine: 60 ( 1 894); Barratte in Bonnet & Barratte,
PI. vase. Tunisie: 73 (1896); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 179 (1925); Stefanoff in God. Agr.-les.
Fak. Univ. Sofiya 11: 175 (1933), 12: 87 (1934), \nPflanzenareale
4( 1 ): Karte 6a ( 1 933); Maire in Bull. Soc. Hist. nat. Afr. N.21:2\5
(1936); Quezel & Santa, Nouv. Fl. Algerie 2: 683 (1963); Pott.-
Alap., Fl. Tunisie 1: 509 (1979). Type: Algeria, prope La Calle,
1785, Poiret s.n. (P-holotype).

Fig. 25E, Map 31.

H. perforatum sensu Poir., Voy. Barbaric 2: 224 (1789).
Icon: Coss., Illfl. Atlant. 2: t. 102 (1892).

Perennial herb ( l-)2 m tall, erect, ± suffrutescent at base, branched
all up stem, with branches short, strict to ascending, glabrous. Stems

STUDIES IN HYPERICUM

vinous red, narrowly 2-lined but otherwise terete, not ancipitous,
internodes exceeding leaves; cortex exfoliating in sheets. Leaves
sessile; lamina 10-15(-20) x 3-5(-ll) mm, narrowly oblong to
elliptic-oblong or lower often obovate to oblanceolate, paler be-
neath, papyraceous, thinly glaucous, plane, spreading to deflexed;
apex rounded, margin recurved and ± undulate, base cuneate to
cordate-amplexicaul; venation: 3(4) pairs of main laterals ascending
from lower 0.25-0.5 of midrib, midrib branched above, tertiary
reticulation obscure; laminar glands pale, dense, unequal, relatively
small, not prominent; intramarginal glands black, dense. Inflores-
cence c. 60-flowered from 3^ nodes with branches from up to 2
nodes below, the whole globose and rather congested to broadly
pyramidal and rather lax, sometimes with additional flowering
branches from further down stem; pedicels 3-4 mm; bracteoles
reduced, linear-lanceolate, sparsely black-glandular-ciliate. Flow-
ers 12-15 mm in diam.; buds ovoid-ellipsoid, rounded. Sepals 3-3.5
x 0.7-1 mm, subequal, slightly basally connate, narrowly ovate-
oblong to linear-lanceolate, acute, glandular-denticulate to -ciliate;
veins 5, not branching or 3, branching; laminar glands pale, shortly
linear to striiform or rarely punctiform; marginal glands black,
small, rather sparse. Petals golden? yellow, tinged red dorsally, 7-8
x 3-3.5 mm, c. 2.3 x sepals, narrowly elliptic to oblong-elliptic,
rounded, with apiculus amber- or black-glandular; laminar glands
pale, linear to distally punctiform; inframarginal glands black,
sparse, on inner margin only. Stamens c. 35-40, longest c. 5.5-6.5
mm, c. 0.8 x petals; anther gland black. Ovary c. 2 x 1 mm, narrowly
ovoid-pyramidal; styles c. 3 mm, 1.5 x ovary, erect-outcurved.
Capsule 5-6 x 3—4 mm, ovoid-pyramidal to ovoid, exceeding se-
pals, usually enclosed by petals twisted together. Seeds
straw-coloured, c. 0.7 mm long; testa finely linear-foveolate.

Marshes, streamsides and damp moors; c. 300-700 m.

NW Tunisia (Mogods, Kroumirie), Algeria (Constantine, E. Al-
giers).

TUNISIA. Mogods: Cap Serrat(/We Pettier- Alapetite, 1979). Kroumirie:
Foret d'A'in-Draham, 15 September 1885 (fr), Robert & Cosson in Soc.
Dauph. (1887) 5205 (BM); a Test de Tabarque [Tabarka], 6 July 1883 (fl),
Cosson et al. s.n. (K); Am Draherru [Draharu], 18 September 1893 (fr),
Robert s.n. (K); Djebel Gorra (fide Pettier- Alapetite, 1979).

ALGERIA. Constantine: prope La Calle, June 1 839 (fl), Bove s.n. (K);
inter Philippeville [Skidka] et Bone [ Annaba], in paludosis ditionis Senhadja,
3 July 1 86 1 (fl), Letourneux & de la Perraudiere in Kralik, PI. Alger. 108 (K);
pres Bone, du versant sud de 1'Eydough [Dj. Eydough], June 1866 (fl & fr),
Trabut in Frag. Fl. alger. exsicc. 562 (BM). Alger: fide Quezel & Santa
(1963).

H. afrum is quite different from other Mediterranean species of
Hypericum. It has been associated with//, elegans Stephan ex Willd.
(sect. 9. Hypericum) by Stefanoff (1933a, 1934) and with //.
undulatum Schousboe ex Willd. in the same section by me (Robson,
1977a); but it does not fit well into sect. Hypericum. On the other
hand, all the characters discordant in that section fit in sect. 27.
Adenosepalum (e.g. the old petals twisting together over the devel-
oping capsule, and the pale linear-foveolate seeds); and//, afrum can
easily be interpreted as the third species in a northward cline: //.
conjungens-H. kiboense-H. afrum. It must therefore be a relict
species of considerable antiquity.

6. Hypericum aethiopicum Thunb., Prodr.fl. cap. : 1 38 ( 1 800), Fl.
cap.: 439 (1823); Choisy in DC., Prodr. 1: 552 (1824); Eklon &
Zeyher, Enum. pl.Afr. austr.: 53 (1835); Sonder in Harv. & Sond.,
Fl. cap. 1: 117 (1860); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 179 (1925); Exell & Mendonca, Consp.
Fl. Angolensis 1: 120 (1937); Bredell in Bothalia 3: 577 & map

179

(1939); Robson in Kew Bull. 3: 438 & map (1958), in Exell &
Wild, Fl. iamb. 1(2): 382, t. 73A (1961); Gibson, Wildfls Natal
(coast, reg.): t. 67, f. 4 (1975); Killick & Robson in J. Ross, Fl.
southn Afr. 22: 17, f. 4 (1976). Type: South Africa, Cape Prov-
ince, 'Crescit in Robbeberg, Houtniquas' (fl), Thunberg s.n. (a
UPS-lectotype, K!-sketch; f3 UPS, Kl-sketch).
Map 32.

Icon: J. Ross, Fl. southn Afr. 22: 18, f. 4 (1976).

Perennial herb sometimes woody at base, 0.1-0.45(-0.6) m tall,
with stems arising from woody rootstock, erect or decumbent,
unbranched, glabrous. Stem green to vinous red, sometimes ancipi-
tous or 2-lined above, otherwise terete, eglandular or
black-gland-dotted, internodes shorter than to exceeding leaves;
cortex sometimes exfoliating in strips. Leaves sessile or rarely to 0.4
mm petiolate, 5-25 x 3-15 mm, ± broadly ovate or more rarely
orbicular or elliptic-ovate to oblong, paler beneath, sometimes
glaucous beneath or on both sides, plane, chartaceous to membra-
nous, spreading or ± appressed; apex obtuse or rarely subacute to
rounded, margin entire, often recurved, base cordate-amplexicaul to
rounded; venation: 4 pairs of laterals curved-ascending from lower
0.4-0.7 of midrib, tertiary reticulation rather lax, often obscure;
laminar glands pale or rarely 1-2 black, dense, unequal, sometimes
prominent; intramarginal glands black, rather dense. Inflorescence
\-c. 25-flowered from up to 3 nodes, curved-corymbiform, lax or ±
congested, occasionally with flowering branches from lower nodes;
pedicels 2-4(-5) mm; bracteoles reduced, ovate to linear-lanceolate
or narrowly elliptic, with dense marginal black glands sometimes
prominent or on cilia. Flowers 15-25 mm in diam.; buds ellipsoid,
acute. Sepals 4-8 x 1.5-2 mm, subequal, lanceolate or linear-
lanceolate to oblong, acute to acuminate, entire to glandular-ciliate;
veins 5, unbranched; laminar glands pale and/or black, striiform to
punctiform; inframarginal or marginal or marginal glands black,
stometimes on cilia, dense. Petals canary yellow, usually tinged red
dorsally, (8-)10-13(-15) x 3-5 mm, 1-4 x sepals, elliptic to
oblanceolate-spathulate, rounded to subtruncate with apiculus obso-
lete, subapical; laminar glands mostly black, ± elongate-punctiform
with a few pale, striiform to punctiform; marginal to inframarginal
glands black, rarely prominent, distally dense. Stamens c. 50-70, ±
indistinctly 3(4)-fascicled, longest 7-10 mm, 0.7-0.8 x petals;
anther gland amber. Ovary 3(4)-locular, 2-4 x 1 .5-2 mm, ovoid;
styles 3(4), 3-6.5 mm, (1.7-)2-3 x ovary, spreading-incurved.
Capsule 6-7 x 3^4 mm, narrowly ovoid-pyramidal to ellipsoid,
shorter than to slightly exceeding sepals, enclosed by petals twisting
together. Seeds chestnut brown, c. 1 mm long; testa finely linear-
foveolate.

180

N.K.B. ROBSON

Map 32 Sect. 27: 6. H. aethiopicum: a. subsp. sonderi
subspecies (arrow) *.

specimens, O records; b. subsp. aethiopicum A specimens, A records; intermediates between

Open grassland or more rarely bare or cultivated areas or seasonal
swamps in high-rainfall areas; 135-1850 m.

South Africa (Cape Province, Natal, Orange Free State, Transvaal),
Lesotho, Swaziland, Zimbabwe (eastern), Mozambique (central),
Angola (Huila).

H. aethiopicum is most closely related to 3. H. conjungens, having a
± herbaceous habit with the woodiness confined to below ground
level or sometimes the base of the stems. It can be divided into two
subspecies, of which the northern (subsp. sonderi} is the less special-
ized in most characters except for the black-gland-dotted stems. The
southern subsp. aethiopicum has eglandular stems, but the sepal
margin is glandular-ciliate, not entire, and the habit is never as robust
as it is in some Transvaal and Natal plants. There is also a northward
reduction trend, the plants of subsp. sonderi from Zimbabwe, Mo-
zambique andAngola being smaller with smaller, usually ± orbicular
leaves. The correlation of gland-dotted stems and entire sepals
occasionally breaks down, hence the reason for making these taxa
subspecies.

6a. Hypericum aethiopicum subsp. sonderi (Bredell) N. Robson
inKewBull. 12:440&map(1958),inExell &Wild, Fl. zamb. 1:
382 (1961); Letty, Wild fls Transvaal: 220, t. 109 f. 3 (1962);
Trauseld, Fl. pis Natal Drakensberg: \ 25 & photographs ( 1 969);
Jacot Guill., Fl. Lesotho: 211 (1971); Killick & Robson in J.
Ross, Fl. southn Afr. 22: 19, f. 4.2 (1976); Fabian & Germish.,

Transvaal wild fls: 178, t. 84d (1982). Type as for H. sonderi
Bredell.

H. aethiopicum var. glaucescens Sond. in Harv. & Sond., Fl. cap. 1:
118 (1860); Burtt Davy, Man. pi. Transvaal: 251 (1926). Type:
Transvaal, Magalisberg, Aapjesrivier, October (fl), Zeyher 149
(S-lectotype, selected here; BM!, LU!). The citation by Sonder
('at Aapjes River and Macallisberg, Zeyher') implies that more
than one Zeyher specimen was seen, and so the above specimen
has been selected as the lectotype. Killick & Robson (1976)
mention Burke s.n. from Magalisberg (BM!, K!) as possibly
another syntype.

H. aethiopicum sensu Sond. in Harv. & Sond., Fl. cap. 1: 118
(1860), pro parte; Baker f. in J. Linn. Soc. Lond. (Bot.) 40: 26
(1911); Eyles in Trans. Roy. Soc. S. Afr. 5: 420 ( 1 9 1 6); Norlindh in
Bot. Notiserl934: 101 (1934); Exell & Mendonca, Consp. Fl.
Angolensis 1(1): 120 (1937), et auct. plur.
H. aethiopicum var. huillense Engl. in Bolm Soc. broteriana 17: 83

(1900), nomen.

H. sonderi Bredell in Bothalia 3: 578 & map ( 1 939); Verdoorn in Fl.
pis S. Afr. 23: t. 8897 (1943). Types: (numerous specimens cited,
from Transvaal to Cape Province) Natal, in planitie prope Cam-
perdown, 900 m, 17 September 1893 (fl), Schlechter 3270
(KMectotype, Robson, 1958; G!, PRE!, Z!-isolectotypes).
H. sonderi var. transvaalense Bredell in Bothalia 3: 579 & map
(1939). Types: Transvaal, Pietersburg Distr., Woodbush, Moun-
tain Home Farm, 1530 m, 18 December 1935 (fl), Mogg 13996

STUDIES IN HYP ERIC UM

181

(PREMectotype, Robson, 1958; K!); Shiluwane, Junod 4290
(PRE!-syntype); without locality, Wager TRV7223 p.p. (PRE!-
syntype).

Icones: Exell & Wild, Fi zamb. 1: 384, t. 73A (1961); J. Ross, Fl.
southnAfr. 22: 18, f. 4.2 (1976).

Stems black-(or occasionally amber-)gland-dotted. Sepals entire or
almost so, with marginal glands immersed or ± prominent but not on
denticles or cilia.

Northern Cape Province (Kokstad) north to eastern Zimbabwe and
central Mozambique; Angola (Huila).

ANGOLA. Huila: Humpata, December (fl), Newton 161 (COI*).

ZIMBABWE. East: Inyanga, prope dejectum fluminis Pungwe, c. 1800
m, 6 November 1930 (fl), Fries, Norlindh & Weimark 2733 (BM, LU, PRE);
Umtali, Stapleford Forest Reserve, Hope Patrol, 1800 m, 16 October 1959
(fl), Chase 7174 (BM, SRGH); Melsetter, Chirinda, 20 October 1947 (fl),
Wild 2070 (K, SRGH).

MOZAMBIQUE. Manica & Sofala: Mossurize, Espungabera, 13 No-
vember 1943 (fl), Torre 6187 (BM, LISJC*).

SWAZILAND. Hlatiku, Verdun, c. 750 m, 14 November 1986 (fl),
Compton 26322 (K, PRE*)11; Siteki-Mhlumeni road, 15 km from Siteki, by
fence round Blue Jay Ranch, 575 m, 29 August 1978 (fl), Prior 187 (K).

TRANSVAAL. Zoutpansberg Distr.: near Entabeni, 19 August 1930
(fl), Hutchinson 4238 (BM, K). Pietersburg Distr.: Duiwelskloof, 9 July 1929
(fl), Galpin 9401 (K, PRE*). Pilgrim's Rest Distr., Pilgrim's Rest, lower end
of town above Velijde R., 18 November 1937 (fr), Galpin 1454 (K, PRE*).
Lydenburg Distr.: bei Lydenburg, January 1884 (fl), Wilms 137 (BM, K, Z).
Pretoria Distr.: Magaliesberg, c. 1400 m, 12 December 1955 (fr), Schlieben
7682 (K, Z). Rustenburg Distr.: c. 2.9 km NE of Derby Stn, 1530 m, 28
January 1962 (fl & fr), Acocks 21971 (K, PRE*). Belfast Distr.: Rictolei,
Crocodile R., June 1932 (fl), Smuts 29 (K). Barberton Distr.: Saddleback
Mtn, southern slopes, 1095-1440 m, October-November 1890 (fl), Galpin
1116 (GRA*, K). Ermelo Distr.: near Sheepmoor, February 1960 (fr), de
Winter 75 19 (K, PRE*).

ORANGE FREE STATE. Ventersdorp Distr.: Ventersdorp, 1530 m, 7
November 1966 (fl), Bayliss 37 10 (Z). Harrismith Distr.: Harrismith, 1 800 m,
November 1904 (fl), Sankey 22 (K). Ladybrand Distr.: Ladybrand, 1650 m,
27 December 1958 (1. fl), Werdermann & Oberdieck 1566 (B*, K).

NATAL. Vryheid Distr.: Twekloof, Altemooi, December-January 1 925-
6 (fl), ThodeM 146 (K). Hlabisa Distr.: Zululand, 330 m, 22 November 1953
(fl), Ward 1758 (K, PRE*). Louwsburg Distr.: Itala Nature Reserve, 1200-
1260 m, 7 December 1975, Brown & Shapiro 13 (K, PRE*). Ladysmith
Distr.: near Cathedral Peak, near Uhlumbonje R., 3 November 1957 (fl),
Goodier 350 (K, SRGH*). Bergville Distr.: Tugela valley, Mont-aux-Sources,
1650 m, 20 February 1926 (fl), Bayer & McClean 212 (K, PRE*).
Pietermaritzberg Distr.: between Drummond and Cato Ridge, January 1974
(fr), Stirton 1008 (K, PRE*). Port Shepstone Distr.: Marina Beach, Strey 5950
(PRE*).

LESOTHO. Lereibe, 1913?(fl),Dfeter/en358(K);22kmfromOuthing,
near Senka bridge, 1520m, 10 December 1 977 (\.fl\Killick 4370 (K, PRE*).

CAPE PROVINCE. Griqualand East: Glen Hope, November 1913 (fl),
Jaconet & Jacottet 570 (Z). ? Distr.: near Fraser Hills, Ntsubane Forest Stn,
1400 m, 24 August 1976 (fl), Venter & Vorster 121 (K, PRE*). Aliwal North
Distr.: Doctors Drift, Gerstner 137 (PRE*).

6b. Hypericum aethiopicum subsp. aethiopicum

Icon: J. Ross, Fl. southnAfr. 22: 18, f. 4.1 (1976).

Stems eglandular or very rarely with a few amber gland dots. Sepals
with margin black-glandular-denticulate or -ciliate distally or wholly.

South-eastern Cape Province (Griqualand East to Riversdale).

CAPE PROVINCE. Griqualand East: without precise locality, February,
Tyson 1376 p.p. (PRE*). Mt Currie Distr.: Kokstad, Glengarry Road near
Kokstad, 12 January 1976 (fl), Coleman 876 (K, NH*). Matatiele Distr.:
Cedarville, November, Bandert 102 (GRA). Herschel Distr.: Sterkspruit,

11 Southern African records with an asterisk (*) have been seen by Killick but not by me.

May, Hepburn 380 (GRA*). Albert Distr.: Albert district, 186- (fl), Cooper
1774 (K). Stutterheim Distr.: Stutterheim Commonage, 780 m, 27 December
1942 (fl), Acocks 9544 (K, PRE*). Cathcart Distr.: Kabousie R., December,
Flanagan 794 (GRA*, SAM*). King William's Town Distr.: King William's
Town, Flanagan 2142 (PRE*, SAM*). Victoria East Distr.: Victoria East,
Rattray 130 (PRE*). Stockenstroom Distr.: Katberg, 24 November 1902 (fl),
Sole 583 (Z). Fort Beaufort Distr.: Adelaide, Great Winterberg, January, Ford
11410 (PRE*). Bedford Distr.: Bedford, December, Binnie 213 (GRA*).
Albany Distr.: Albany, Highlands Station, 1800 m, 1 June 1978 (fl), Bayliss
8794 (Z). Somerset East Distr.: in clivis ad latera Montis Boschberg prope
Somerset East, c. 900 m, December 1 874 (fl), MacOwan 397 (K). Alexandria
Distr.: Karlsrand, near Karl Landsman Memorial, 360 m, 9 November 1954
(fl), Johnson 1085 (K). Uitenhage Distr.: ad mentis radices 'Chumiberg'
(Kafferland), September-October 1 835 (fr), Ecklon 4 1 5 (FR, K); Port Eliza-
beth, Zuurberg Sanatorium, 1 November 1930(fl),L0Hg225(K). Humansdorp
Distr.: Assegaibos, November, Marloth 1093 1 (PRE*). Knysna Distr.: Prince
Alfred's Pass, 900 m, 15 December 1937 (fr). Wall 368 (LU); below
Boukamma & Ruigte Vlei, 135 m, April 1928 (fl), Fourcade 3919 (K).
George Distr.: Kamanasie Hills, near George, November 1847 (fl). Prior s.n.
(K, PRE*). Riversdale Distr.: Corente R., Muir 855 (PRE*, SAM*).

Coleman 846 (from Kokstad) is one of the specimens mentioned
above as being morphologically intermediate between the two sub-
species. It comes from the narrow zone of overlap between the
respective areas of the subspecies, as does Jacottet & Jacottet 570
(Griqualand, Glen Hope (Z)). Bredell's (1939) record of subsp.
aethiopicum from Herschel district, near the Lesotho border (Map
32), may be an error or an intermediate form. I have not seen the
specimen (Hepburn 380 (GRA)).

Subsect. 2. Pubescentes N. Robson in Bull. nat. Hist. Mus.
Land. (Bot.) 23: 69 (1993). Type: H. pubescens Boiss. (see
p. 90).

Perennial herbs, with indumentum up to sepals or lower part of
inflorescence or rarely only to base of inflorescence; leaves free;
bracts and bracteoles not glandular-auriculate. Species 7-12.

7. Hypericum pubescens Boiss., Elench. pi. nov.: 26 (1838), Voy.
hot. Espagne 1: t. 36, 2: 1 15 (1840); Walp., Repert. hot. syst. 1:
383 ( 1 840); Ball in/ Linn. Soc. London 16: 374 ( 1 877);Amo, Fl.
fan. Penins. Ibericat: 24 (1878); Willk. & Lange, Prodr. fl.
hispan. 3: 592 (1878), Suppl.: 272 (1893); Batt. & Trab., Fl.
Algerie, Dicots. 1: 183 (1888); Maire in Mem. Soc. Sci. nat.
Maroc no. 7: 180 (1924); Stefanoff in God. Agr.-les. Fak. Univ.
Sofiya 11: 175 (1933), 12: 87 (1934), in Pflanzenareale 4(1):
Karte 6a (1933); Sennen & Mauricio, Cat. fl. Rif orient.: 26
( 1 934); N. Robson in Tutin et al., Fl. europaea 2: 266 ( 1 968); Ali
in AH & Jafri, Fl. Libya Guttiferae: 8, f. 4 (1976); Haslam, Sell &
Wolseley, Fl. Maltese Is.: 198 (1977); Pignatti, Fl. Italia 1: 347 &
map (1982); Ramos in Trab. Depto Bot. Univ. Complut. 12: 54, t.
6f. 1 (19%2\ \r\Actabot.Malacit. 11: 169,f.9b(1986), inValdes,
Talavera & Fern.-Gonz., Fl. vase. Andalucia occ. 1: 3 1 8 & fig. &
map (1987); Burdet, Charpin & Jacquemoud in Candollea 39:
789 (1984); Greuter, Burdet & Long, Med-Checklist 3: 273
(1986); Ramos in Castroviejo et al., Fl. iberica 3: 184 (1993).
Type: Spain, Granada, Sierra Nevada, inter Estepona et San
Roque,Boissiers.n. (G-lectotype, Ramos, 5 December 1980; see
Burdet et al., 1984).

Fig. 26A, Map 33.

H. tomentosum subsp. pubescens (Boiss.) Ball in J. Linn. Soc.
London (Bot.) 16: 374 (1877); Jahand. & Maire, Cat. pis Maroc
2: 485 (1932), Suppl.: 1071 (1941); Quezel & Santa, Nouv. Fl.
Algerie 2: 682 (1963); Pott.-Alap., Fl. Tunisit 509 (1979).

H. tomentosum [van] y pubescens (Boiss.) Perez Lara in An. Soc.

182

N.K.B. ROBSON

Map 33 Sect. 27: 7. H. pubescens specimens •, probably this species
but recorded as H. tomentosum by Pignatti (1982) O; 8. H. psilophytum

Esp. Hist. nat. 24: 333 (1895); Fiori, Nuov. Fl. Italia 1: 525 (1924).
H. aegusanum Tineo ex Lojac., Fl. sicul. 3: 431 (1908), nomen.
H. tomentosum subsp. pubescens var. viridulum Pau in Bol. R. Soc.

esp. Hist. nat. 22: 57 (1922); Jah. & Maire, Cat. pis Maroc 2: 485

(1932). Type: Morocco, Xauen, VldalyLopezs.n.(MA-ho\otype).
H. tomentosum subsp. pubescens var. damnatorum Maire in Bull.

Soc. Hist. nat.Afr. N. 29: 41 1 (1938). Type: Algeria, hot springs of

Hammam-es-Skoutin (Aquae Tibelitanae), Maire s.n. (AL-

holotype).
H. tomentosum sensu Guss., Fl. sicul. syn. 2(1): 28 (1843); E.

Durand & Barratte, Fl. libic. Prodr.: 48 (1910); R. Keller in Engl.

& Prantl, Nat. Pflanzenfam. 2nd ed. 21: 179 (1925); Borg, Descr.

fl. Maltese Is.: 249 (1927) et auct. plur, pro parte omnes excl.

typum.

Icon: Boiss., Voy. hot. Espagne 1: t. 36 (1840).

Perennial herb c. 0. 1-0.7 m tall, erect to decumbent or ascending or
rarely prostrate, from ± woody taproot, branching, rooting and
sometimes woody at base, upper branches sometimes present, spread-
ing to ascending, the whole plant up to the sepals (dorsally)
greyish-villous to -pubescent. Stems green to pale reddish, 2^-lined
and compressed in and near inflorescence, otherwise terete,
internodes mostly exceeding leaves. Leaves sessile; lamina 6-40(-
70) x 2-16(-23) mm, narrowly oblong or oblong-lanceolate to
rarely ovate-oblong, concolorous, chartaceous, not glaucous, plane,
spreading; apex subobtuse to usually rounded, margin plane, entire
or very rarely lower leaves sparsely glandular-subdentate, base
rounded to usually cordate-amplexicaul; venation: 3(4) pairs of
laterals curved-ascending from base or up to lower 0.4 of midrib,
tertiary reticulation rather dense, often obscure; laminar glands pale,
dense, subequal, not prominent, sometimes also with a few black,
punctiform, scattered; intramarginal glands black, rather dense to
sparse; marginal glands (very rarely present) amber, globose, promi-
nent. Inflorescence (l)3-c. 5-flowered from up to 3 nodes,
curved-corymbiform, becoming monochasial after first flower, some-
times with flowering branches from up to most lower nodes, the
whole then broadly triangular to cylindric; pedicels 1.5-3.5 mm;
bracts not auriculate; bracteoles linear, usually with a few sessile
black glands or (proximally) black-glandular-cilia. Flowers 15-30
mm in diam.; buds cylindric-ellipsoid, obtuse. Sepals 5-10 x 1.5-
2.5 mm, subequal, linear-lanceolate to lanceolate, ± long-aristate,
entire to subentire; veins 3(5), unbranched; laminar glands pale,
punctiform to slightly elongate; marginal glands black, 3-5 on each
side, sessile, the arista reddish and very rarely with small black
apical gland. Petals pale to bright yellow, often veined dorsally, 9-15
x 3-6.5 mm,c. 1 .5 x sepals, oblanceolate, rounded, apiculus apiculate
or absent; laminar glands pale, striiform to punctiform; marginal to
inframarginal glands black, not or scarcely prominent. Stamens c.
30-50, clearly 3-fascicled, longest 6-1 1 mm, 0.6-0.65 x petals;
anther gland black. Ovary 2-3 x 1-2 mm, ovoid-ellipsoid to ellip-
soid; styles 3, 5-7 mm, 2-3 x ovary, widely spreading-incurved.
Capsule 6-7 x 3.5-5 mm, ovoid, shorter than sepals, enclosed by
petals twisted together. Seeds purplish brown, 0.6-1 mm long; testa
finely scalariform. 2n=18 (Robson, 1981), 36 (fide Ramos, 1987:
318);Reynaud, 1986.

River and stream margins, rocky ground, roadsides and waste habi-
tats, often in damp ground; 0-1875 m.

Portugal (south), Spain (south), Morocco, Algeria, Tunisia, Malta,
Sicily, Sardinia? (see below).

PORTUGAL. Algarve (fide Franco, Nova Fl. Portugal 1 : 450 ( 1 97 1 )).

SPAIN. Huelva and Sevilla: bothfide Valdes et al., Fl. Vase. Andal. Occ.
1: 318 (1987). Cadiz: Puerto Santa Maria, Pinal del Cato, 8 June 1849 (fl),
Bourgeau 104 (BM); 1 0 km from Rota, by Naval Base, 13 May \979(f\),P.F.
Cannon 454 (RNG). Granada: prope Granatum, 1 849 (fl), Boissier & Renter
s.n. (K). Murcia: Elche, July 1902 (fl & fr), Prah & Sylva s.n. (BM). Valencia:
all specimens from this province seen so far appear to be (hybrid) intermedi-
ates (see p. 184). Catalonia: Catalogne, Hostaleb (?), June 1906 (fl & fr),
Herb. Lacaita 19056 (BM).

GIBRALTAR. Gibraltar, n.d. (fl.), Gay s.n. (BM).

MOROCCO. Tanger: ad pedem montis Zem-Zem (El Haus), 4 m, 12
June 1930(fl), Font Quer 425 (BM, Z); halfway between Tangier and Tetuan,
250 m, 2 July 1973 (fl & fr), Davis 54714 (BM, E*). Rif: prope Ein-Zeren
(Beni Uriaguel), 30 m, 20 July 1928 (fl & fr), Font Quer 306 (BM, Z);
Targuist to Chechaouen, 40 km E. of Chechaouen, 28 June 1 974 (fl), Reading
U./Brit. Mus. Exped. 1098 (BM). Oranais littoral: Martimprey-du-Kiss a
Aghbal, 13 June 1931 (fl), Faure s.n. (K). Centre nord: Volubilis, ruines
romaines, 500 m, 3 June 1978 (fl), Lewalle 8854 (BM, BR*); 16 km from El-
Hajab to Fes, 26 June 1974 (fl), Reading U./Brit. Mus. Exped. 1065 (BM).

STUDIES IN HYPERICUM

183

Fig. 26 A. H. pubescens: (a) habit; (b) leaf; (c) elongating inflorescence; (d) sepal; (e) petal; (0 capsule with two enclosing petals. B. H. tomentosum: (g)
habit; (h) flower bud. C. H. collenettiae: (i) habit; (j) flower bud; (k) petal. D. H. sinaicum: (1) habit; (m) flower bud (a, c, g, i, 1 x '/z; b x 1; d, f, h, j, k, m
x 3). A. Davis 54993. B. Davis 59519. C. Collenette 3752. D. (1) Schimper 42 1 , (m) Collenette 7589.

184

N.K.B. ROBSON

Monts de Za'i'an: 30 km above Azrou, road to Ifrane, 1300 m, 26 June 1974
(fl), Reading U./Brit. Mus. Exped. 1052 (BM). Nord-ouest: c. 10 km SSE of
Rabat, valley of Oued Akrech, 22 May 1 96 1 (fl), J. & P. de Wilde & Dorgelo
2384A(BM,WAG*);propeoppidumRabat,3May \926(f\),Lindberg 1661
(H, K). Sud-ouest: Cap Blanc au sud-ouest de Mayagan, 15 June 1950 (fl),
Sauvage 8537 (K); Cap Beddouza [Meddouza], N. of lighthouse, 30 m, 1
June 1974 (fl), Reading U./ Brit. Mus. Exped. 116 (BM). Sud steppique:
Sheshoua, May 1871 (fl). Hooker s.n. (K); Merrakish, Tanga, 30 June 1984
(fl), Lewalle 1 1022 (BM, BR*). Sud-ouest littoral: Cap Sari, 8 May 1926 (fl),
Lindberg 1846 (H); environs d'Agadir, Sidi-Moussa, 21 May 1877 (1. fl),
Ibrahim s.n. (K). Plaine de Sous: Taroudannt, 12 June 1920? (fl), Lynes 8
(BM). Moyen Atlas: Khenifra, village Imam Ines, 1300m, 25 June 1981 (fl),
Lewalle 9961 (BM, BR*); Ladla, Beni Mellal, 500 m, 28 May 1925 (fl),
Jahandiez 379 (K). Grand Atlas: Jebel Sarhro, SE of Boumalne du Dades,
July 1969(fl), fiwvfcr 1 8 (BM); Tizi-n-Test to Taroudannt, 1100m, 12 June
1974 (fl), Reading U./Brit. Mus. Exped. 570 (BM). Anti Atlas: 5 km from
Tafraoute to Igherm, 1 100 m, 9 June 1974 (fl), Reading UJBrit. Mus. Exped.
451 (BM); 2 km from Igherm to Taroudannt, 1800 m, 10 June 1974 (fl),
Reading U./Brit. Mus. Exped. 554 (BM).

ALGERIA. Oran: Oued Imbert, 4 August 1929 (fl), Faure s.n. (BM);
plaine du Lac Sale, July 1849 (fl), Roman s.n. (BM); Sidi Bel-Abbes to
Tlemcen, between Ben Badis and Oued Mimoun,c-. 1300m, 7 June 1975 (fl),
Davis 58818 (BM. E*). Alger: Blidah, Oued-el-Kebir, lOAugust 1864 (1. fl),
Lefebvre s.n. (K). Kabylie: between Les Falaises andTichi (Bejaia-Djidjelli),
s.l., 28 May 1 97 1 (fl), Davis 52932 (BM, E*); Kerrata, 800 m, July 1 897 (fl),
Reverchon 9 (BM). Constantine: Constantine, in declivitate humidis montis
Mansourah, 6 July 1 869 (fl), Paris 314 (BM); Constantine, July 1 876 (fl & e.
fr), Reboud 1 144 (FR). Hauts-Plateaux oranais: Ravin de Tafaroua, 10 July
1868 (fl), Allariou (K). Atlas Saharien constantinofs: M'chouneche prope
Biskra, 3 June 1902 (fl), Chevallier PI. Sah. Alg. 416 (K).

TUNISIA. Kroumerie, Nord-est and Cap Bon: fide Pottier-Alapetite
(1979: 509). Dorsale: Zaghouan, 26 July 1854 (fl), Kralik 196 (BM, K).
Centrale: sud de Sousa [Sousse], 8 June 1883 (fl), Cosson et al. s.n. (K).

LIBYA. Tripolitania: Misurata, 22April 1939 (fl), Simpson 39662 (BM),
SandwithZ7l9(K).

MALTA. Malta: Corradino, Wied Kerdu, April-June 1929 (fl), Reade
s.n. (BM); Qaliet, 21 May 1965 (fl), Llanfranco s.n. (BM).

SICILY. Trapani: Trapani, July 1908 (fl), Albo s.n. (BM). Palermo:
Palermo, in campis maritimis, 1902 (fl & fr), Ross s.n. (BM).

SARDINIA (extinct?). Nuoro: prope Laconi, 1 827 (fl), Muller s.n. (H)
(see opposite).

CULTIVATED. England: London, Chelsea Physic Garden, 1767 (fl),
Herb. Chelsea 2272 (BM).

H. pubescens is directly related to the Canary Island 1. H.
glandulosum and appears to be ancestral to 8. H. tomentosum. Its
distribution is south-west of that of H. tomentosum, but there are
quite wide areas of overlap in south Spain, north-eastern Morocco
and north-western Algeria. Morphological intermediates are few
and apparently mainly confined to two small regions, viz. Valencia
near L. Albufera and the Moroccan-Algerian border. In addition, the
recorded chromosome numbers (H. pubescens 2n=18, 36; H.
tomentosum 2n=16) suggest that they are specifically distinct. For
these reasons it seems best to treat the intermediates as hybrids and
H. pubescens and H. tomentosum as species.

Although H. pubescens is quite variable in size and habit, depend-
ing on humidity and exposure, at least one of the named variants is
not worthy of recognition: var. damnatorum was based on the
occurrence of purplish anthers, which was probably due to suffusion
of hypericin from the anther gland. I have not seen the type of var.
viridulum

7x. Hypericum pubescens x tomentosum

Map 34.

H. tomentosum var. intermedium Coss. [in Bourgeau, Exsicc. 1852:
no. 1582 (1852) nomen] ex Willk. & Lange, Prodr.fl. hispan. 3:
592 (1878); Sagredo, FL Almeria: 290 (1987). Types: Spain,

Valencia, ad forsa interViveret Jerica, August 1850(fl), Willkomm
Iter hisp. II 478 (BM!); in sabulosis pineti prope lacum Albufera,
August 1850 (fl), Willkomm Iter hisp. II 506 (BM!); ad lacum
Albufera, 1852, Cosson in Bourgeau Exsicc. 1852 No. 1582 (P-
lectotype, selected here; K!).
H. tomentosum [var.]agenuinum subvar. elevatum Perez Lara in An.

Soc. esp. Hist. nat. 24: 333 (1895), nom. illegit. superfl. Type as

for H. tomentosum var. intermedium Coss.
?//. tomentosum [var.] (3 ambiguum Perez Lara in An. Soc. esp. Hist.

nat. 24: 333 ( 1 895). Type' : Spain, Andalucfa, Gades, Perez Lara.

No specimen; described from living plants.

Intermediate in form between the parents. Hair length as in H.
pubescens. Inflorescence variable but usually nearer that of H.
tomentosum. Sepals either narrowly lanceolate with small gland on
the arista or broadly lanceolate to elliptic without apical gland, but
always with marginal glandular cilia.

Spain (Valencia, Gerona), Morocco (Rif, Oranais littoral), Algeria
(western Oran).

SPAIN. Valencia: Lac Albufera, July, Graels s.n. (BM). Gerona: Llers,
14 July 1907 (fl), Sennen 273 (FR, H, RNG).

MOROCCO. Rif: Hidum, 10 May 1930 (fl), Sennen & Mauricio sub
7542 (BM). (One sheet as above, the other (BM) unaltered asTafersit, 28 June
1930 (fl), Sennen & Mauricio 7542). Oranais littoral: Martimprey-du-Kiss, a
Aghbal,fl. 13June 1931, Faure s.n. (BM). Moyen Atlas: Oulad AH, 1950m,
30 July 1975 (fl), Stones 28050 (BM).

ALGERIA. Oran: prope Tlemcen, 18 June 1856 (fl), Bourgeau s.n.
(BM).

In addition, the only Sardinian specimen of H. tomentosum seen
(q.v.) tends towards H. pubescens, and it seems probable that the
populations of that species from southern Italy do likewise.

8. Hypericum psilophytum (Diels) Maire in Bull. Soc. Hist. nat.
Afr.Nord26: 195 (1935), op. cit. 31: 14(1940); Jahand. & Maire,
Cat. pis Maroc, Suppl.: 1070 (1941); Quezel & Santa, Nouv. Fl.
Algerie, 2: 682 (1963) ['psilophyton']; Greuter, Burdet & Long,
Med-Checklist 3: 273 ( 1 986). Type: Algeria, Ahaggar Hochland,
Seitental des Tig'ameiin-en-tisita, 24 April 1914 (fl), Geyr 111
(Bt-holotype).

Map 33.

H. suberosum var. psilophvtum Diels in Bot. Jb. 54, Beibl. 1 20: 100

(1917).

H. tomentosum subsp. psilophytum (Diels) Maire in Mem. Soc. Hist,
nat. Afr. Nord 3: 154 (1933); Ozenda, Fl. Sahara 2nd ed.: 342, f.
117(1977).

Icon: Ozenda, Fl. Sahara 2nd ed.: f. 1 17 (1977).

Perennial herb 0.1-0.5 m tall, tufted, decumbent to ascending from
woody taproot, branching at base, upper branches few or absent,
stems spreading- to appressed-puberulous, leaves papilliform-
puberulous, inflorescence glabrous or almost so. Stems becoming
purplish red, 2-lined near inflorescence, otherwise terete, internodes
shorter than to exceeding leaves. Leaves sessile; lamina 5-1 3 x 2.5-

STUDIES IN HYPER1CUM

185

6 mm, ± narrowly oblong to oblanceolate, concolorous, chartaceous,
not glaucous, plane, suberect; apex rounded, margin plane, entire,
base rounded to subcordate-amplexicaul; venation: 2 pairs of laterals
curved-ascending from lower 0.4 of midrib, tertiary reticulation
obscure; laminar glands pale, rather dense, subequal, not prominent,
with a few black, punctiform, scattered; intramarginal glands black,
rather dense; marginal glands absent. Inflorescence 2-c. 25-flow-
ered, from 1(2) nodes, curved-corymbose, monochasial after first
flower or sometimes wholly monochasial, sometimes with 1-2
branches; pedicels 1.5-3 mm; bracteoles linear, black-glandular-
ciliate. Flowers c. 15 mm in diam., stellate; buds ellipsoid-
subglobose, subacute. Sepals 2-4 x 1-1 .2 mm, lanceolate or elliptic
to narrowly oblong, subequal, acute to shortly aristate, margin
glandular-ciliate; veins 3, unbranched; laminar glands pale, shortly
linear to punctiform, sometimes also black, punctiform; marginal
glands black, sessile or on short cilia, apex glandular. Petals yellow,
tinged red dorsally, 7-8 x c. 3 mm, 2 x sepals, oblanceolate to
narrowly elliptic-oblong?, rounded, apiculus absent; laminar glands
pale, striiform to punctiform; marginal glands black (?). Stamens c.
15-25, 3?-fascicled, longest 4-5 mm, c. 0.65 x petals; anther gland
black. Ovary c. 2 x 1 mm, narrowly ovoid; styles 3, c. 2.5-3 mm,
1.25-1.5 x ovary, spreading (-incurved?). Capsule 3.5-4 x 2.5 mm,
ovoid to ovoid-subglobose, 1.5-2 x sepals. Seeds not seen; testa
striate (fide Ozenda). 2n=18 (Reynaud, 1986).

Damp margins of wadis; c. 800-2040 m.

Morocco (GrandAtlas, western Sahara), Algeria (Teffedest, Ahaggar,
Tassili n'Ajjer).

MOROCCO. Grand Atlas: Gorges du Todha [Todrha], 1 500 m, 23 June
1939 (fl), Maire & Weiller473 (AL*, P). Maroc desertique occidental: Djebel
Bani aTatta, 8 April 1934 (o. fr), Maire & Wilczek It. Maroc. XXIV s.n. (AL*,

P).

ALGERIA. Sahara septentrional. Teffedest:^ deQuezel & Santa ( 1 963).
Ahaggar: Tazerouk, 1950 m, 1928, Maire 262 (AL*); Oued Tessekimt, 2040
m, 1928, Maire 262 (AL*); ibid., 1400-1500 m, 1928, Maire 263 (AL*).
Tassili n'Ajjer: valley of OuedTidjamain-n-Tisita, 800-900 m, 29 April 1914
(fl), Geyr 272 (Bf).

H. psilophytum is most closely related to 7. H. pubescens rather than
to 9. H. tomentosum, the relationship favoured by Maire (1933). In
particular it is near the reduced forms of//, pubescens from the Anti-
Atlas of Morocco. It is not far geographically, morphologically or
ecologically from these to the Moroccan populations of //.
psilophytum, which occur on the south-western parts of the Jebel
Bani and Grand Atlas, next to the Sahara.

H. psilophytum appears to be a good species, differing from //.
pubescens in several characters including the shorter, mostly
appressed indumentum, the smaller, sometimes glandular-ciliate
sepals and the glabrous or slightly puberulous inflorescence. From
the literature the (isolated) Algerian populations would seem to be
more distinct than the Moroccan ones.

9. Hypericum tomentosum L., Sp. pi.: 786 (1753); Willd., Sp. pi.
3: 1466 (1802); Choisy, Prodr. monogr. Hyperic.: 52 (1821), in
DC., Prodr. 1: 551 (1924); Moris, FL Sardoa 1: 322, t. 21 (1837)
pro parte excl. ref. Decaisn.; Rchb., lc.fl. germ. helv. 6: t. 346, f.
5183 (1844); Ball in/ Linn. Soc. London (Bot.) 16: 374 (1877),
pro parte quoad spec, ex Tangier, Willk. & Lange, Prodr. fl.
hispan. 3: 592 (1878), SuppL: 111 (1893); Batt. & Trab., Fl.
Algerie, Dicots. 1: 183 (1888); Bonnet & Baratte, Expl. sci.
Tunisie, Cat. pi: 74 (1896); R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed 21: 179 (1925), pro parte; Cout., Fl.
Portugal 2nd ed.: 483 (1930); Stefanoff in God. Agr.-les. Fak.
Univ. Sofiya 10: 58, t. 4 f. 22 (1932), 11: 174 (1933), pro parte

excl. distr. ital. merid. et Sicilia et Melita, 12: 87 (1934), in
Pflanzenareale 4( 1 ): Karte 6a (1933); Jahand. & Maire, Cat. Pis
Maroc 5: 485 (1934); Sennen & Mauricio, Cat.fl. Rif oriental: 26
(1934); Samp., Fl. Portugueza 2nd ed.: 323 (1946); N. Robson in
Tutin et al., Fl. europaea 2: 266 (1968); Franco, Nova Fl. Portu-
gal 1: 450 (1971); Fournier, Quatre Flores de France 2nd ed.:
456 (1977); Barcelo, Fl. Mallorca 3: 178 (1979, reprint 1990);
Pott.-Alap., Fl Tunisie: 509 (1979), pro parte quoad subsp. eu-
tomentosum Maire; Pignatti, Fl. Italia 1: 347 (1983); Ramos in
Trab. Depto Bot. Univ. Complut 12: 54, t. 6 f. 1 (1982); Greuter,
Burdet & Long, Med-Checklist 3: 273 ( 1 986); Ramos inActa hot.
Malacit. 11: 169,f. 9a(1986), inValdes,Talavera&Fern.-Gonz.,
Fl. vase. Andalucia occ. 1: 317 (1987), in Castroviejo et al., Fl.
iberica 3: 1 84 (1993); N. Robson in Cullen et al., Eur. Gdn Fl. 4:
72 ( 1 995). Type: The Linnaean phrase-name clearly describes //.
tomentosum rather than H. pubescens Boiss. Following Clusius
and Bauhin, Linnaeus differentiated a smaller French plant (H.
supinum tomentosum minus & monspeliacum of Bauhin) from a
larger Spanish one (H. supinum tomentosum majus & hispanicum
of Bauhin), the latter as a var. (3. From Clusius (Rar. pi hist. 2:
181, 1601)it is clear that these are both forms of//, tomentosum.
There are three specimens of this species that Linnaeus studied,
two in his own herbarium (LINN 943.42 and 43) and one in Herb.
Burser (UPS 16: 23). Specimen 943.42 is labelled Hypericum
supinum majus hispanicum and was given to Linnaeus by
Monnier; 943.43 is a typical young-flowered plant of the north-
ern, French form, the collector being unknown; and the Burser
plant is a fruiting specimen from Montpellier. Of the two speci-
mens of the French form, I therefore select the flowering one
(LINN 943.43) as the lectotype of H. tomentosum L.
Fig. 26B, Map 34.

H. lusitanicum Poir., Encycl., Suppl. 3: 702 (1814); Choisy, Prodr.

monogr. Hyperic.: 57 (1821), in DC., Prodr. 1: 553 (1824); Willk.

& Lange, Prodr. fl. hispan. 3: 592 (1878); Debeaux in Act. Soc.

Linn. Bordeaux 42: 1 64 ( 1 889); R. Keller in Engl. & Prantl, Nat.

Pflanzenfam. 2nd ed. 21: 180 (1925); Stefanoff in God. Agr.-les.

Fak. Univ. Sofiya 10: 58, t. 1 f. 16 (1932), 11: 175 (1933), 12: 87

(1934), in Pflanzenareale 4(1): Karte 6a (1933); Samp., Fl.

Portugueza 2nd ed.: 323 (1946). Type: Portugal, Anon, in Herb.

Desfontaines (P-lectotype, selected here); Hypericum

tomentosum, lusitanicum, minimum Tournefort, Inst. R. Herb.:

256(1 700) (P-syntype).

H. suberosum Salzm. ex Boiss., Voy. hot. Espagne 2: 1 15 (1839) in
synon.; Bonnet & Barratte, Expl. sci. Tunisie, Cat. pi.: 74 (1896),
nomen. 'Type': Morocco, Tangier, Salzmann s.n. (C).

Map 34 Sect. 27: 9. H. tomentosum specimens •, Tunisian records A
(Pettier- Alapetite, 1979); 7. H. pubescens x 9. H. tomentosum O.

186

N.K.B. ROBSON

H. supinum Vis. in Atti Riun. Sci. ital. : 1 75 ( 1 84 1 ), ///. piante Grec.
Asia minors: 17 (1842); N. Robson in Notes R.B.G. Edinb. 27:
196 (1967); Mabberley in Taxon 31: 71 (1982).Types: (i) as for//.
tomentosum L. [van] p, Sp. pi. 2nd ed.: 1 106 (1763) - lectotype,
Robson (1967«); (ii) Turkey, circa Antandro ad sinum Golfo
d'Adramitti dictum 1819, Parolini & Webb (PAD!-syntype).
Robson (1967a) showed that de Visiani's clear intention was to
raise Linnaeus's H. tomentosum [var.] (3 to specific rank and to
include the Turkish specimen (from Edremit) in that species. The
Turkish specimen belongs to 21 . //. atomarium, but the lectotype
must be that of H. tomentosum [van] (3 L., i.e. LINN 943.42 (see
discussion under H. tomentosum).

H. tomentosum var. dissitiflorum Roemer in Linnaea 9: 17 (1852);
Willk. & Lange, Prodr. Fl. hisp. 3: 592 ( 1 878); Cout., Fl. Portugal
2nd ed.: 483 (1939). Type: Spain, Jaen, Sierra Morena, inter
Aldeaquemada et S. Esteban, Willkomm (COI-holotype).

?//. canescensTrevir., Hyper, sp. animadv. : 1 0 ( 1 861 ). Type: Stefanoff
(1934: 87) made this a synonym of H. lusitanicum Poiret, and
Treviranus suggested that it might be a prostrate form of that
species. He stated, however, that the fruit [of the type specimen,
now in BHU?] had been destroyed. The identity of this taxon must
therefore remain subjudice.

Adenosepalum tomentosum (L.) Fourr. inAnn. Soc. Linn. Lyon N.S.,
16:352(1868).

H. tomentosum var. glabrescens Porta in Nouv. Giorn. hot. ital. 19:
301 (1887), nomen. 'Type': Balearic Is., Mallorca, prope pagum
Arta, 10-20 m, 30 June 1885 (fl), Porta & Rigo s.n. (BM!).

H. tomentosum var. racemosum Batt. in Batt. & Trab., Fl. Algerie,
Dicots.: 183 (1888); Cadevall Gotes, Fl. Catalunya 1(4): 379
(1915). Type: Algeria, ? (AL).

H. tomentosum var. palustre Batt. in Batt. & Trab., Fl. Algerie,
Dicots.: 183 (1888). Types: Algeria (Boufarik, Maison-Carree,
etc.) (AL-syntypes).

H. tomentosum subsp. lusitanicum (Poir.) Willk. in Willd. & Lange,
Prodr. fl. hispan., Suppl.: 272 (1893).

H. tomentosum [var.] a genuinum Perez Lara in An. Soc. esp. Hist,
not. 24: 332 (1895). Type as for H. tomentosum L.

H. tomentosum [var.] y lusitanicum (Poir.) Perez Lara in An. Soc.
esp. Hist. nat. 24: 333 (1895).

H. tomentosum subsp. eu-tomentosum Maire, Cat. Pis Maroc 2: 485
(1932). Type as for H. tomentosum L.

H. tomentosum subsp. carbonelli Sennen & Mauricio, Cat. fl. Rif
orient.: 26 (1934), nomen.

H. carbonelli Sennen & Mauricio [Campagn. hot.: 105 (1936) in
obs., nomen] Diagn. nouv.: 185 (1936). Type: Morocco, Rif,
Tarquist, Bab-Izugar, 1 926, Emberger & Maire s.n. (BC-holotype).

H. tomentosum subsp. eu-tomentosum var. carbonelli (Sennen &
Mauricio) Maire in Bull. Soc. Hist. nat. Afr. Nord 27: 216 (1936).
Type as for H. carbonelli.

H. tomentosum subsp. wallianum Maire in Bull. Soc. Hist. nat. Afr.
Nord 27: 79 (1936), in torn, cit.: 216 (1936); Jahand. & Maire,
Cat. Pis Maroc, Suppl.: 1071 (1941). Type: Morocco, Grande
Atlas oriental pres de Rich, May (fl), Wall (AL?-holotype).

H. tomentosum var. densifolium Sennen (description not traced).
Type: Spain, Murcia, Sierra de Espuna, Goto de Sta Eulalia, 21
June 1927 (fl), Jeronimo in Sennen, Pis d'Espagne 6419 (BC-
holotype; BM!-isotype).

H. tomentosum var. ramosissimum Sennen (description not traced).
Type: Spain, Murcia, Sierra de Espuna a Sta Eulalia, 480 m, 8 July
1929 (fl), Sennen & Jeronimo s.n. (BC-holotype; BM!-isotype).

Icones: Rchb., Ic. fl. germ. helv. 6: t. 346 (1844); Bonnier, Fl. ill.
France 2: t. 104(1912).

Perennial herb c. 0.09-0.53 m tall, erect or decumbent to prostrate,
from scarcely woody taproot, branching and sometimes rooting at
base, upper branches sometimes present in some or all axils, spread-
ing to ascending, the whole plant up to sepals (dorsally) greyish
villous to -tomentose or leaves hirsute to crisped-pubescent. Stems
green, terete, internodes mostly exceeding leaves. Leaves sessile;
lamina 5-26 x 2-1 1 mm, elliptic-oblong to oblong or oblanceolate
or ovate to triangular-ovate, concolorous, chartaceous, not glaucous,
plane, spreading; apex rounded, margin plane, entire, base cuneate
to truncate or subcordate; venation: 3 pairs of laterals curved-
ascending from lower 0.3-0.4 of midrib, tertiary reticulation obscure;
laminar glands pale, dense, unequal, not prominent; intramarginal
glands black, rather dense to sparse, sometimes irregular and be-
coming submarginal, or absent (then black glands wholly absent).
Inflorescence 3-c. 70-flowered from up to 3 nodes, curved-
corymbiform to cylindric, with flowering branches from up to most
lower nodes, the whole then cylindric; pedicels 0.5-2 mm, elongat-
ing to 5-8 mm in fruit; bracts not auriculate, bracteoles linear, with
apex and margin black-glandular-ciliate. Flowers 10-15(-20) mm
in diam.; buds ellipsoid, obtuse to rounded-obtuse. Sepals 3-5(-6?)
x 1-2.5 mm, subequal to unequal, lanceolate to ovate or broadly
elliptic, acute to usually shortly aristate, with marginal glands
mostly prominent; veins 3-5, unbranched; laminar glands pale,
punctiform to slightly elongate, sometimes also 1-3 black, submar-
ginal; marginal glands black, c. 8-16 on each side, on cilia or sessile
but then nearly always prominent, the apex usually with larger
gland. Petals bright yellow, not tinged or veined red, 6-1 1 x 2.5-
3.5(^1?) mm, c. 2 x sepals, oblanceolate, rounded, apiculus lateral,
shortly acute to absent; laminar glands pale, proximally linear,
distally striiform to punctiform; marginal to inframarginal glands
black, few, subterminal, not prominent. Stamens c. 25-35, clearly 3-
fascicled, longest 5-7 mm, 0.6-0.8 x petals; anther gland black.
Ovary 1 .5-2 x 1-1.3 mm, narrowly ovoid-pyramidal; styles 3, 5-5.5
mm, 2.75-3.3 x ovary, widely spreading-incurved. Capsule 4-5 x
3-3.5 mm, ovoid-subglobose, shorter than sepals, enclosed by petals
twisting together. Seeds 'greyish-brown' (fide Ramos, 1987), c. 0.8
mm long; testa finely reticulate-scalariform (fide Ramos, 1983: 57,
t. 6 f. 1). 2n = 16 (Queiros, 1991), n = 8 (Nielsen, 1924).

STUDIES IN HYPER1CUM

Stream margins, damp or marshy grassland, maquis, evergreen
scrub; 0-200 m (France), 0-800 m (Italy), 0-1200 m (Spain), 0-
1450 m (Morocco).

Portugal (N. to Coimbra), Spain (except NW), France (Mediterra-
nean lowlands), Italy (Liguria, also Campagnia and Basilicata /ufe
Pignatti, 1 982), Corsica (one locality, extinct?,// tfcGamisans, 1 985),
Sardinia, Majorca, Morocco (Tangier, Rif, northern Moyen Atlas),
Algeria (north), Tunisia (north).

PORTUGAL. Coimbra: Estrada Coimbra-Cantanhede, entre Sao
Fagundo e Anca, 20 September 1956 (fl), A. & R. Fernandes & Matos 61 20
(BM, COP). Leiria: Caldas da Rainha, 2 July 1889, Murray s.n. (BM).
Santarem: Estrada Abrantes-Sardoal a 3 km de Sardoal, 18 June 1956(fl),/4.
& R. Fernandes & Santos 5837 (BM, COI*). Lisboa: Estoril district, 1939
(fl), Ogilvie 18 (K). Setubal: ad pedem Serrae da Arrabida, June 1842 (fl),
Welwitsch 189 (K); Arrabida, April 1987 (st), Bowen 5018 (RNG).

GIBRALTAR. Near Gibraltar, Campomento, May 1868 (fl), Hirst 23
(BM).

SPAIN. Cadiz: 2 km S. of El Bosque, 400 m, 12 July 1981 (fl), M. & S.
Gardner 1 126 (BM, RNG). Sevilla: between Carmona and Seville, 20 June
1 927 (fl), Wilmott s.n. (BM). Malaga: Cartama, 2 July 1 888 (fl), Reverchon 69
(BM, K); c. 4 km NE of Archidona, 10 June 1964 (e. fl), Sandwith 6312 (K).
Granada: Nevada ad pedem Mt Domago, 22 July 1 873 (fl), Wmklers.n. (BM).
Jaen: Sierra de Cazorla, Cueva del Polvo, 1050-1200 m, 28 July 1951 (fl),
Heywood 1772 (BM). Albacete: Sierra de Segura, 15 km W. ofYeste, 1.5 km
E. of Prados, 1000 m, 30 June 1979 (fl), Reading UJBot. Dept. Exped. 563
(BM, RNG). Murcia: Sierra de Espufia, Goto de Sta Eulalia, 21 June 1927
(fl), Jeronimo in Sennen 641 9 (BM). Valencia: Dos Aguas, 15 May 1984 (fl),
Peris & Stubing Exsicc. II 48 (K, RNG). Cuenca: Serrania de Cuenca, 1 6 July
1967 (fl), Rivas-Goday & Morja (RNG, MAF*). Valladolid: Olmedo, 28
June 1851-2 (fl), Lange s.n. (K). Burgos: Miranda, 30 July 1909 (fl), Elias
806 (BM). Navarra: environs de Lumbier, entre Artieda et Ripodas, c. 500 m,
July 1972 (fl), Vivant \r\AuquierE\s., Fasc. 15, 6722 (H).Teruel: Valacloche,
800 m, July 1 893 (fl), Reverchon 626 (BM, FR). Barcelona: Castelloglels, 1 2
June 1 920 (fl), Sennen s.n. (K). Gerona: Llers, 1 August 1 907 (fl), Sennen 422
(FR, H, RNG).

FRANCE. Herault: Murviel les Montpellier, 14 July 1895 (fl & fr),
Mandon s.n. (BM). Bouches du Rhone: Meyreuil, entre le pont de Bayou et
Beaureceuil, 27 June 1 9 1 2 (fl), Delmas 1 9 (BM). Vaucluse: Avignon, Rignien,
n.d. (fl & fr), Bentham s.n. (K).Var: Le Cannet-des-Maures, 10 July 1950 (fl),
Bouchard in Exsicc. B. de Retz fasc. 5, 1 656 (K). Alpes Maritimes: Nice a St
Andre, June 1894 (fl), Bonafons s.n. (BM).

ITALY. Liguria: Bordighera, Val di Sasso, 3 July 1 890 (fl), Bicknell s.n.
(BM, K). Also recorded from Campagnia and Basilicata (see opposite).

CORSICA. Near Bonifacio, Pouzolz 'in Hb. Eor.\fide Briquet (1935).

SARDINIA. Nuoro: ad rivulis prope Laconi, July 1827 (fl), Muller s.n.
(H, K).This specimen shows characters verging towards//, pubescens (see p.
184).

BALEARIC ISLANDS. Majorca: Puigpunyent, c. 1.5 km S. of Puig-
punyent towards Galilea, 340 m, 14 July 1969 (fl), L & I. Ferguson 2491
(BM); Alcudia, between Mai Pas and Barcarets, 9 August 1987 (fl & fr), J. &
M. Cannon 3363 (BM); Puerto Pollensa, near sea level, 27 November 1934
(fl & fr), Martindall 79 (K).

MOROCCO. Tanger: Tanger to Sebta (Ceuta), 5 km, 70 m, 1 8 June 1 987
(fl), Jury, Rejdali & Watson 830 1 (BM, RNG); Tetouan, 450 m, 2 1 June 1 977
(fl), Lewalle 8762 (BM, BR*). Rif: between Ketama and Taounate, 134 km
from Fez, 1300-1400 m, 8 July 1973 (fl), Davis 54967 (BM); Atlas Rifain:
Beni-Bufrah, 1300 m, 19 June 1934 (fl), Sennen & Mauricio 9642 (BM).

ALGERIA. Alger: between Boufarik and Kolea, c. 40 m, 28 June 1 975
(fl), Davis 59519 (BM); gorges de la Chiffa, vers de Camp des Chenes, 12
June 1875 (fl), Cosson s.n. (K).

TUNISIA. No specimens seen. Recorded from north and down eastern
plain to Sidi el Hani and Mahdia by Pottier-Alapetite (1979).

H. tomentosum tends to be smaller in all its parts than 7. H.
pubescens and to have shorter hairs; but (i) the south Moroccan
populations of H. pubescens are vegetatively smaller than many
examples of//, tomentosum, although the flowers are still relatively
large, and (ii) the populations of H. tomentosum from the Atlas

187

Rifain of northern Morocco described as H. carbonelli have an
unusually long indumentum. None of the described variations in H.
tomentosum, however, appears to merit taxonomic recognition. For
intermediates between H. tomentosum and H. pubescens, see Sp. 7x
(p. 184).

The reasons for treating H. tomentosum and H. pubescens as
distinct species are discussed on p. 1 84. H. tomentosum has a mainly
west Mediterranean distribution, from NE Morocco and NW Alge-
ria to Liguria with Majorca and Corsica; H. pubescens has a
south-west Mediterranean distribution, south Morocco to SE Spain,
Tunisia, Malta and Sicily. Where these distributions meet (Sardinia)
or overlap (Morocco-Algeria border, SE Spain), there are intermedi-
ates (hybrids). The records from southern Italy would therefore be
expected to be of//, pubescens, but they are in fact of//, tomentosum
(Pignatti, 1982). Pignatti (in litt. 1991) has informed me that the
Basilicata record (from Monte Vultura) is based on an old specimen
of Terraciano (in Fl?), which I have yet to see. The record(s) of H.
tomentosum from Campania are similarly doubtful, as are those
from Tunisia (Pottier-Alapetite, 1979).

1 0. Hypericum somaliense N. Robson inKew Bull. 13: 396 ( 1 959);
Cufod. in Bull. Jard. hot. Etat. Brux. 29, Suppl.: 589 (1959),
Moggi & Pisacchi in Webbia 2: Til, f. 14, carta 6 (1967). Type:
Somalia (North): Markat, 1440m, 1 2 August 1957 (ft), Newbould
886 (K!-holotype).

Map 35.

Icon: Moggi & Pisacchi in Webbia 22: 279, f. 14 (1967).

o

Perennial herb 0. 1 5-0.6 m tall, erect or decumbent to ascending and
rooting at base, from ± woody taproot, upper branches sometimes
present, ascending, the whole plant up to the bracteoles whitish

Map 35 Sect. 27: 10. H. somaliense •;!!.//. collenettiae D; 12. H.

188

tomentose. Stems green to pale red-brown, terete, internodes mostly
exceeding leaves. Leaves sessile; lamina 10-18(-23) x 4-9 mm,
narrowly oblong to triangular-ovate or triangular-lanceolate, almost
concolorous, chartaceous, not glaucous, plane, spreading; apex
obtuse to apiculate or rounded, margin plane, entire, base rounded to
subcordate-amplexicaul; venation: 2-3 pairs of laterals curved-
ascending from lower 0.2-0.4 of midrib, tertiary reticulation obscure;
laminar glands pale, dense, unequal, not prominent; intramarginal
glands black, rather dense. Inflorescence c. 10-30-floweredfrom 1-
2 nodes, subcorymbiform to capitate-rounded, rather dense, with
sparse amber glands, not prominent, without lower flowering
branches; pedicels 2.5^.5 mm in fruit; bracts not auriculate;
bracteoles linear-triangular, with margin black-glandular-ciliate.
Flowers c. 15 mm in diam.; buds ellipsoid, acute. Sepals 2-5 x 1.5-
2 mm, equal, lanceolate, to ovate, acute to aristate, glandular-ciliate
to -fimbriate; veins 5(-7), unbranched; laminar glands pale, linear to
punctiform; marginal glands black; marginal and apical glands
black, smaller. Petals orange-yellow, not red-tinged, 7-9 x c. 2-3
mm, c. 2-3.5 x sepals, oblong-lanceolate?, rounded, with apiculus
acute, short; laminar glands pale, linear to striiform; marginal glands
black, sessile, ± prominent. Stamens c. 30, clearly 3-fascicled,
longest 6.5-8 mm, c. 0.9 x petals; anther gland black. Ovary c. 2 x
1 mm, ovoid-conic; styles 3, c. 6 mm, c. 3 x ovary, spreading-
incurved. Capsule 4.5-5.5 x 2.5-3 mm, narrowly ovoid, exceeding
sepals, not enclosed by separately twisting petals. Seeds dark red-
dish brown, 0.5 mm long; testa finely scalariform-reticulate.

Streamsides, clay flushes; 1440 m.

Somalia (North).

SOMALIA. Ragad, 1440 m, 12 August 1957 (fl), Newbould 895 (K).

Only the above two collections have been made of H. somaliense,
which thus appears to be a relict species of very restricted distribu-
tion. Its relationships are clearly with 7. H. pubescens, from which
it differs by the dense, mostly glabrous inflorescence and the glandu-
lar-ciliate to -fimbriate sepals with a row of submarginal black
glands.

1 1 . Hypericum collenettiae N. Robson in Bull. nat. Hist. Mm.
Land. (Bot.) 23: 69 (1993). Type: Saudi Arabia, Asir, Taif-Abha
road 82 km S. of Baljurshi, Wadi Mahra, 1 800 m, 5 August 1982
(fr), Collenette 3752 (BM!-holotype; K!-isotype).

Fig. 26C, Map 35.

H. sp. aff. sinaicum sensu Collen., ///. guide fls Saudi Arabia: 262 &
photographs (1985).

Icon: Collen. (1985), see above.

Perennial herb or subshrub, c. 0.15-0.2 m tall, erect from woody

N.K.B. ROBSON

taproot or decumbent at base and rooting, often with short ascending
branches below inflorescence, the whole plant up to the inflores-
cence ± densely white-puberulous. Stems green to purplish red,
terete, internodes mostly shorter than leaves. Leaves sessile, mark-
edly tetrastichous; lamina 7.5-c. 13x1 .5-4 mm, narrowly oblong to
(upper) triangular-lanceolate, concolorous, chartaceous, appearing
greyish blue but not glaucous, plane, spreading; apex obtuse, margin
plane, entire, base rounded to subcordate; venation: 2 pairs of
laterals curved-ascending from lower c. 0.25 of midrib, tertiary ret-
iculation not seen; laminar glands pale, dense, unequal, not
prominent; intramarginal to submarginal glands black, rather sparse.
Inflorescence (l-)5-c. 11 -flowered, from 1(2) nodes, sub-
corymbiform to rounded-pyramidal when flowers numerous, rather
lax, without lower flowering branches; pedicels 3-7 mm in fruit,
with ± prominent reddish glands; bracts not auriculate; bracteoles
linear-lanceolate, with margin black-glandular-ciliate. Flowers c. 25
mm in diam.; buds ellipsoid, acute. Sepals 3.5-4 x 0.8-1.2 mm,
equal, ± narrowly lanceolate, acute to subaristate, irregularly glan-
dular-ciliate or with prominent sessile glands; veins 5, outer branched;
laminar glands pale, linear to striiform and (toward margins) black,
punctiform to striiform; marginal and apical glands black, smaller.
Petals golden yellow, not red-tinged, 9-10(7-12) x c. 4.5-5 mm, c.
2.5 x sepals, oblong-obovate, truncate, with apiculus obtuse, short;
laminar glands distal, pale, shortly striiform to punctiform and
black, ± elongate-punctiform, toward apex, marginal or inframar-
ginal glands black, few, elongate-punctiform. Stamens c. 40, clearly
3-fascicled, longest 5-6 mm, c. 0.6 x petals; anther gland black.
Ovary c. 2 x 1 .5 mm, ovoid-pyramidal; styles 3, c. 4 mm long, c. 2
x ovary, widely spreading. Capsule 4.5-5 x 3-4- mm, ovoid-cylin-
dric, exceeding sepals, enclosed by petals twisting together. Seeds
dark red-brown, 0.7 mm long; testa finely scalariform-reticulate.

Shady rock crevices; c. 1 800 m.

Saudi Arabia (Asir).

SAUDI ARABIA. Asir: Taif-Abha road, 70 km S. of Baljurshi, Wadi
Mahra, 1829 m, 16 April 1979 (fl), Collenette 1401 (K); Bashwat [between
Baha and Abha], 9 August 1975 (1. fl), A. El-Sheikh in Herb. KSUH 1067
(KSUH-photograph).

H. collenettiae is intermediate geographically and in most respects
morphologically between 10.//. somaliense and 12. H. sinaicum. As
in H. somaliense, the stems are erect and apparently not rooting; the
leaves and indumentum are nearer those of H. sinaicum, as is the
relatively lax and few-flowered inflorescence. In having black laminar
petal-glands it also resembles H. sinaicum, but in its red-glandular
pedicels it is nearer to H. somaliense. The prominence of these
glands, however, distinguishes it from both species.

H. collenettiae is known as yet from only two populations and is
therefore, like H. somaliense, a relict species. According to Mrs
Collenette (verb, comm.), the Baljurshi population may no longer
exist.

12. Hypericum sinaicum Hochst. [in Steud.,Nomencl. hot. 2nd ed.
1: 789 (1840), nomen] ex Boiss., Fl. orient. 1: 808 (1867); R.
Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 179
( 1 925); Post, Fl. Syria 2nd. ed., 1: 233 ( 1 932); Stefanoff in God.
Agr.-les. Fak. Univ. Sofiya 10: 58, t. 3 f. 29 (1932), 11: 163
(1933), 12: 85 (1934), \nPflanzenareale 4(1): Karte 4a (1933);
Montasir & Hassib, ///. man. Fl. Egypt 1: 309 (1956); Tackh.,
Students' Fl. Egypt 2nd ed. 1: 150, t. 42B (1974); Greuter,
Burdet & Long, Med.-Checkl. 3: 273 ( 1986). Type: Egypt, Sinai,
'in rupibus ad scaturigines reg. Bestan ad rad. m. Sinai', 3 July
1835 (fl & fr), Schimper42\ (G-holotype; BASBG!, E!, JE!, K!,
LE!-isotypes), sphalm 42 (Boiss.).

STUDIES IN HYPERICUM
Fig. 26D, Map 35.

H. tomentosum sensu Decne. in Annls Sci. nat. (Bot.) II, 3: 287

(1835).

Icon: Tackh., Students' Fl. Egypt 2nd ed. 1: 151, t. 42B (1974).

Perennial herb 0.1-0.35 m tall or long, ascending to prostrate from
woody taproot, with ± numerous near-basal branches sometimes
rooting, otherwise unbranched below inflorescence, the whole plant
except pedicels and flowers densely to sparsely and shortly whitish
pubescent. Stems green to reddish, terete, internodes exceeding
leaves. Leaves sessile or with petiole toe. 0.3 mm; lamina 6- 18 x 3-
8 mm, narrowly oblong to elliptic or (lower) oblanceolate, almost
concolorous, thinly chartaceous, not glaucous, plane, spreading to
ascending; apex obtuse to rounded, margin plane, entire, base
broadly cuneate to subcordate; venation: 2-3 pairs of laterals curved-
ascending from lower 0.3-0.6 of midrib, tertiary reticulation obscure;
laminar glands pale, rather dense, unequal, not prominent, and
black, few, scattered, especially towards apex; intramarginal glands
black, rather sparse, or absent; submarginal glands black, sparse, not
always distinguishable from black laminar ones. Inflorescence c. 3-
20-flowered from 1-2(3) nodes, corymbiform to rounded-pyramidal;
pedicels 2.5-3.5(^4-) mm in fruit; bracts not auriculate; bracteoles
lanceolate or oblong to linear, with apex and margin sparsely black-
glandular-ciliate or some glands sessile. Flowers 9-1 3 mm in diam.;
buds broadly ellipsoid, obtuse to rounded. Sepals 3-3.5 x 1-2 mm,
subequal, ovate or ovate-elliptic to suborbicular, acute to obtuse,
with marginal glands dense to sparse, on cilia or sessile and ±
prominent; veins 5, unbranched; laminar glands pale, linear or long
striiform to punctiform, and a few black, submarginal and occasion-
ally truly laminar; marginal and apical glands black, c. 8-12 on each
side. Petals pale yellow, sometimes veined red, (5-)8-10 x (1 .7-)2-
3 mm, 2-3 x sepals, oblong-lanceolate, rounded, apiculus absent;
laminar glands distal, pale to reddish, striiform to punctiform, and
sometimes a few black, punctiform; marginal glands black, distal,
sessile or on short cilia. Stamens 25-30, clearly 3-fascicled, longest
4-6.5 mm, 0.55-0.8 x petals; anther gland black. Ovary 1.5-2 x c.
1 mm, ovoid-pyramidal to ovoid; styles 3, 2.5^1.5 mm, 1.7-2.2 x
ovary, spreading. Capsule 4-5 x 2-3 mm, ovoid, exceeding sepals,
enclosed by petals twisting together. Seeds dark reddish brown, c.
0.5 mm long; testa finely scalariform-reticulate.

Damp rocks and seepage areas; 900-2400 m.

Egypt (Sinai), Saudi Arabia (extreme N. Hijaz), Jordan (Edom).

EGYPT. Sinai: June 1 832 (fl), Bove \ 52 (E, K); n.d. (fl), Aucher 868 (K);
Mt Sinai, May 1868 (o. fr), Lord s.n. (K); n.d. (fl), Kaiser 115, 288, 536 (Z);
Jebel Katarina, 1800-2400 m, September 1945 (fr), Lord Kinross s.n. (E).

SAUDI ARABIA. Hijaz: Wadi Lauz [Lawz] area near Aqaba, 91 km

189

NE of Al Bud, 1500 m, 8 July 1991 (fr), Collenette 7840 (K); Jabal Lauz,
Wadi Lakus, 900 m, 3 August 1989 (fl), Collenette 7231 (BM, E, K); Jabal
Lauz area near Wadi Abyad, S. of Ha, 21 May 1990 (fl), Collenette 7589
(E*, K).

H. sinaicum is close to 11. H. collenettiae from Asir, but is less
woody at the base, with stems ascending to prostrate (not erect),
leaves broader (sometimes petiolate) and flowers smaller with shorter
and broader sepals. In addition, the pedicels lack the prominent
reddish glands found in H. collenettiae.

Collenette 7231 has delicate slender stems, leaves sometimes
shortly petiolate and relatively small flowers and broad sepals with
sessile marginal glands. Otherwise it seems to fit well into H.
sinaicum, its abnormalities being the result of its growing in a
particularly wet habitat.

Subsect. 3. Caprifolia N. Robson in Bull. nat. Hist. Mus.
Lond. (Bot.) 23: 69 (1993). Type: H. caprifolium Boiss.
(see p. 90).

Subshrubs or usually perennial herbs with indumentum to base of
inflorescence; leaves all or mostly connate in pairs; bracts and
bracteoles glandular-auriculate or not. Species 13-15.

13. Hypericum coadunatum Chr. Sm. in Buch, Phys. Beschr.
Canar.lns.: 153 (1828); Webb & Berth., Phytogr. canar. l:45,t.
4 (1836); Pitard & Proust, Les lies Canaries: 132 (1909); R.
Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed., 2: 180
(1925); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: 58, t. 3
f. 31 (1932), 11: 163 (1933), 12: 84 (1934), in Pflanzenareale
4(1): Karte 4a (1933); Lid in Skr. norske Vidensk.-A.cad. I,
Math.-Nat. Kl., N.S. no. 23: 120 (1967); Bramwell, D. & Z.,
Wild fls Canary Is: 68, f. 41 (1974); Kunkel, Endemismos
Canaries: 292 & map (1977). Type: Canary Islands, Chr. Smith
s.n. (G?-holotype).

Fig. 27B, Maps 36, 37.

H. coadunatum var. disjunctum R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 180, nomen.

Icones: Webb & Berth., Phytogr. canar. 1: t. 4 (1836); Bramwell, D.
& Z., Wild fls Canary Is: f. 41 (1974).

Subshrub or perennial herb with ± woody base, c. 0.75-2 m tall,
erect to ascending from creeping and rooting base with woody
taproot, the branches numerous, forming dense clumps, upper
branches spreading-ascending, the whole plant except inflorescence
sparsely to rather densely crisped-pubescent. Stems green to red-
dish, 2-lined and ancipitous in inflorescence, otherwise terete;
internodes mostly shorter than leaves. Leaves sessile, lower pairs
free, upper pairs connate; lamina 20-40(-48) x 1 5-30(-36) mm,
broadly elliptic to ovate-suborbicular, concolorous (pale green),
herbaceous, not glaucous, plane, spreading; apex rounded, margin

190

N.K.B. ROBSON

Map 36 Sect. 27: 13. H. coadunatum
O (extinct in localities outlined).

(see also Map 37); 14. H. naudinianum D; 15. H. caprifolium A. Sect. 28. 1. H. elodes specimens •, records

plane, base cordate- to subcordate-amplexicaul (lower) or pairs
united with rounded sinus between; venation: 3-4 pairs of laterals,
curved-ascending from lower c. 0.2 of midrib; tertiary reticulation
dense, with major areoles bullate above, impressed beneath; laminar
glands pale, dense near margin but absent near centre, subequal,
sometimes prominent; intramarginal glands black, locally dense but
irregular to sparse or almost absent. Inflorescence c. 1 0-60-flowered
from 1-2 nodes, densely curved-corymbiform, sometimes with
flowering branches from up to 3(?4) nodes, the whole then laxly
cylindric to obconic; pedicels 4-6 mm; bracts not auriculate;
bracteoles linear-triangular, black-glandular-ciliate or with some
marginal glands sessile. Flowers c. 15 mm in diam.; buds ellipsoid,
obtuse. Sepals 3.5-5 x 1.1-1.8 mm, subequal, oblong to lanceolate
or elliptic, acute, with margin gland-fringed; veins 5, branched and
reticulating towards margin and distally; laminar glands pale,
striiform to punctiform and sometimes a few black, subpunctiform;
marginal glands black, prominent. Petals bright yellow, not tinged
red, (6.5-)7-8.5 x c. 2-3.25 mm, c. 1 .6-2 x sepals, narrowly elliptic
to narrowly oblong, rounded, without apiculus; laminar glands pale,
striiform to punctiform; marginal glands black, 1-c. 3, apical, or
absent? Stamens c. 40, clearly 3-fascicled, longest c. 6-7 mm, 0.8-
0.85 x petals; anther gland amber. Ovary 2.3-2.7 x 1 .5 mm, narrowly
ovoid-ellipsoid; styles c. 4.5 mm, c. 1 .6 x ovary, spreading-incurved.
Capsule 4-6 x 3-4 mm, ovoid-subglobose, slightly exceeding se-
pals, enclosed by petals twisting together. Seeds yellow-brown, 0.6
mm long; testa finely scalariform. 2n= 1 8 (Ortega & Navarro, 1 978).

Wet rocks in shade or full sun; 500-1500 m.

Canary Islands (Gran Canada).

CANARY ISLANDS. Gran Canada: San Bartolome de Tirajana, 1000

m, 3 January 1974 (fr), Lewalle 7387 (BM); Cueva del Corcho, 1200 m, 22
July 1972 (fl), Melville & Bramwell 72/26 (K); in aquaeductuum stillicidium,
1 834? (fl), Webb s.n. (K).

Since Maire (1924) wrote that he could find no difference between
the Canarian H. coadunatum and the North African H.
naudinianum Coss. & Durieu, except possibly in the distribution of
gland-dots, most authors have united these species under the earlier
name, H. coadunatum, A detailed comparison, however, has re-
vealed several apparently constant differences betwen them. Thus,
H. coadunatum is more woody in habit with relatively broader
bullate leaves, the lower ones being apparently always free (not all
pairs connate, as in H. naudinianum). It therefore seems desirable to
treat them as separate species. They do, however, have the same
chromosome number (2n=18), which is also found in their nearest
(ancestral) relative, 7. H. pubescens.

Kunkel (1977) reports that H. coadunatum is in danger where it
grows near roads, and that, without protection, it could disappear on
account of lack of water. Keller's ( 1 925) record of ' var. disjunctum"
from Tenerife is apparently without foundation.

14. Hypericum naudinianum Coss. & Durieu in Bull. Soc. hot.
France 2: 308 (1855); Batt. & Trab., Fl. Algerie, Dicots. 1: 182
(1888); Coss.,///. Fl. Atlanticae 2: 10, t. 101 ff. 1-13 (1892); R.
Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 180
(1925); Maire in Bull. Soc. Hist. nat. Afr. Nord 22: 285 (1931).
Type: Algeria, prope Blidah ad amnem Oued-el-Kebir, June
1847, Naudin (P-holotype).

Fig. 27A, Map 36.

H. perfoliatum Munby in Bull. Soc. Bot. France 2: 283 (1855), non
L. (1767).

STUDIES IN HYPERICLJM

191

Fig. 27 A. H. naudinianum: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) capsule. B. H. coadunatum: (f) leaf; (g) sepal. C. H. caprifolium: (h) flower; (i)
sepal. D. H. elodes: (j) habit; (k) leaf; (1) sepal; (m) petal, showing ligule; (n) stamens and ovary, showing one 'lodicule'; (o) flower with developing
capsule, two petals removed (a, j x Vr, b, f, k x 1; h x 2; c-e, g, i, l-o x 4). A. Newbould 186. B. Lewalle 7387. C. Heywood & Davis 408. D. Melderis 87.

192

N.K.B. ROBSON

H. atlanticum Coss. in Bull. Soc. hot. France 22: 56 (1875); R.
Keller in Engl. & Prantl, Nat, Pflanzenfam. 2nd ed. 21: 180
(1925); nomen. Type: Cosson cites Morocco, Djebel Ouensa,
August 1873 (fl), Ibrahim s.n. (K!), but nowhere describes this
'species'.

H. coadunatum var. atlanticum Ball in J. Linn. Soc. (Bot.) 16: 374
(1877); Batt. & Trab., FL Algerie, Dicots. 1: 183 (1888). Type:
Morocco, Grand Atlas, 'in convalle Ait Mesan, versus 1800 m',
13-16 May 1871 (st), Ball s.n. (K!-holotype). Ball's query ('var.
atlanticum nob.?') relates to the Ibrahim collection cited above, of
which a specimen was sent to Ball by Cosson labelled '//.
atlanticum Coss. MSS.'

H. coadunatum sensu Maire in Mem. Soc. Sci. nat. Maroc no. 7: 1 80
(1924); Jahand. & Maire, Cat. pis Maroc 2: 484 (1932); Greuter,
Burdet & Long, Med-Checklist 3: 265 ( 1 986) et auct. afr. plur. pro
parte omnes excl. typum.

H. caprifolium subsp. naudinianum (Coss. & Durieu) Maire in
Jahand. & Maire, Cat. pis Maroc 2: 484 (1932); Quezel & Santa,
Nouvelle Fl. Algerie 2: 681 (1963).

Icon: Coss., ///. FL Atlanticae 2: t. 101, ff. 10-13 (1892).

Perennial herb (0.3-)0.5-1.5 m tall, erect to ascending from creep-
ing and rooting base, with woody taproot, the basal branches
numerous, diffuse, upper branches few, ascending or spreading-
ascending or usually absent, the whole plant except inflorescence
rather densely crisped-pubescent or (leaves) puberulous. Stems green
to reddish (below), sometimes 2-lined and ancipitous in upper parts
of inflorescence, otherwise terete; internodes mostly exceeding
leaves. Leaves sessile, all pairs connate; lamina 10-45 x 10-30 mm,
± broadly elliptic to broadly oblong or obovate, subconcolorous,
herbaceous, not glaucous, plane, spreading; apex rounded, margin
plane, base usually with acute to obtuse or rarely rounded sinus
between pairs; venation: 3 pairs of laterals, curved-ascending from
lower c. 0.3 of midrib; tertiary reticulation dense, plane, not bullate/
impressed; laminar glands pale, dense near margin, sparse near
centre, not prominent and occasionally a few black, scattered;
intramarginal glands black, sparse or absent. Inflorescence c. 10-
100-flowered from 1-3 nodes, densely curved-corymbiform to
rounded-pyramidal, becoming monochasial after second flower,
sometimes with flowering branches from up to 4 nodes, the whole
then laxly cylindric to obconic; pedicels (2-)3-6 mm; bracts not
auriculate; bracteoles linear-triangular, black-glandular-ciliate or
with some or all marginal glands sessile. Flowers c. 15-17 mm in
diam.; buds ellipsoid, obtuse. Sepals 3-5 x 1-1.7 mm, subequal to
unequal, elliptic to oblong or rarely ovate, acute to subacuminate,
with margin gland-fringed; veins 5, branched and reticulating to-
wards margin and distally; laminar glands pale and/or black, striiform
to (mostly) punctiform, scattered; marginal glands black, prominent

or on very short cilia. Petals 'clear butter yellow', tinged or veined
red, 8-9(-10) x c. 2-2.5 mm, narrowly oblanceolate, rounded,
without apiculus; laminar glands pale, punctiform to shortly linear,
very sparse or absent, and occasionally black, few, distal; marginal
glands black, few or in short dense row, distal. Stamens c. 35-40,
clearly 3-fascicled, longest (6-)7-8 mm, c. 0.8 x petals; anther gland
amber. Ovary c. 2.5 x 1.8 mm, narrowly ovoid; styles 3, 4.5-5 mm
long, c. 2 x ovary, widely spreading-incurved. Capsule 4.5-6 x 2.5-
4 mm, ovoid-pyramidal to ovoid, 1.3-2 x sepals, enclosed by petals
twisting together. Seeds yellow-brown, 0.4-0.5 mm long; testa
finely scalariform. 2n = 18 (Jones in Robson, 1981, as H.
coadunatum), n = 9 (Galland, 1988, as H. coadunatum).

Wet rocks, wet shady banks, streamsides, waterfalls; 90-2230 m.

Morocco (Grand Atlas to Rabat Rif), Algeria (north central).

MOROCCO. Nord-ouest: Foret de la Mamora, between Sidi Allal-
Bahraoui and Kenitra, 150 m, 12 April 1971 (st), Davis 54395 (BM, E).
Grand Atlas: Asni,Imlil, 1600m, 18 September 1981 (fl & fr), Lewalle 10037
(BM, BR*); 83 km from Ouazagate, 120 km from Marrakech, along P3 1 c. 9
km S. of Tizi-n-Tickka, 2150 m, 10 July 1987 (fl), Jury, Rejdali & Watson
9223(BM,RNG).Rif:propeAdeldal,BeniSelman,ad 1260m, 12July 1930
(fl), Font Quer 426 (BM). Plaine de Sous: S. of Tizi-n-Test, 10 September
1964 (fl & fr), R. & A. Harley 464 (BM).

ALGERIA. Alger: Belida [Blida], ad rupes madidas secus torrentum
Oued-el-Kebir, 2 July 1864 (fl), Paris 37 (BM); Gorge de la Chiffa, 18 July
1854 (fl), Cosson s.n. (K). Kabylie: recorded from Djurdjura Mts by Quezel
& Santa (1963: 681).

Although very similar to 13. H. coadunatum (q.v.), H. naudinianum
has apparently constant morphological differences as well as its
distinct geographical distribution that favour its recognition as a
species. It is even closer morphologically than is H. coadunatum to
the Spanish 15. H. caprifolium (q.v.), which differs essentially from
it only in the characters of the calyx.

15. Hypericum caprifolium Boiss., Elench. pi. nov.: 26 (1838),
Voy. hot. Espagne 1: t. 35,2: 1 15 (1839);Walp.,/te/*>rr. hot. syst.
1: 383 (1840); Coss., Notes pi. crit.: 99 (1851); Amo, Fl. Iber. 6:
32 (1878); Willk. & Lange, Prodr. fl. hispan. 3: 591 (1878);
Coss., ///. Fl. Atlanticae 2: t. 101 ff. 14-17 (1892); R. Keller in
Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 180 (1925);
Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: 58, t. 3 f. 30
(1932), 11: 163 (1933), 12: 84 (1934), in Pflanzenareale 4(1):
Karte 4a (193); Ramos in Trab. Dep. Bot. Univ. Complut 12: 54
(1983); Burdet, Charpin & Jacquemoud in Candollea 39: 788
(1984); Greuter, Burdet & Long, Med-Checklist 3: 265 (1986);
Ramos inActa hot. Malacit. 11: 169, f. 9d (1986); Motero-Mesa
& Perez-Raya, Fl. Sierra Nevada: 183 (1987); Ramos in
Castroviejo et al., Fl. iberica 3: 182, t. 52 (1993). Type: Spain,
Granada, Sierra Nevada ( 'Sa. Nevada valles'), July 1 837, Boissier
s.n. (G-lectotype, A. Ramos, 1980); loc. cit. ('In montis humidis
regni granatensis praecipue in parte inferiore Sierra Nevada'),
600-1200 m, 1837, Renter s.n. (G-syntype).

Fig. 27C, Map 36.

H. hirsutum Asso, Syn. Stirp. Arag.: 107 (1779), non L. (1753).

Icones: Boiss., Voy. Bot. Espagne 1: t. 35 (1839); Coss., ///. Fl.
Atlanticae 2: t. 101 ff. 14-17 (1892).

Perennial herb 0.2-1 m tall, erect or ascending from creeping and
rooting base, with ± woody taproot, basal branches few, diffuse,
upper branches few, curved-ascending, or usually absent, rarely
branched all down stem, the whole plant except inflorescence ±
densely crisped-pubescent or (some leaves) puberulous. Stems red-
dish, wholly terete or sometimes (2)4(6)-lined in upper parts of

STUDIES IN HYPER1CUM

inflorescence; internodes mostly exceeding leaves. Leaves sessile,
all pairs except lowermost ones connate; lamina (15-)20-50 x (9-)
12-24 mm, elliptic to oblong or ovate-oblong, subconcolorous, not
glaucous, plane, spreading; apex rounded to obtuse, margin plane,
base with sinus between pairs acute to obtuse or (in lowermost)
cuneate; venation: 3 pairs of lateral curved-ascending from lower
0.25-0.5 of midrib; tertiary reticulation dense, plane not bullate/
impressed; laminar glands pale, fairly dense, scattered, small, not
prominent and sometimes black, sparse, scattered; intramarginal
glands black, rather dense, irregular. Inflorescence c. 10-80-flow-
ered from 1-4 nodes, densely curved-corymbiform to
rounded-pyramidal, becoming monochasial after first flower, some-
times with flowering branches from up to 4 nodes, the whole then
laxly cylindric; pedicels 3-5 mm; bracts and bracteoles linear-
triangular, apex aristate, black-glandular-fimbriate, at least
bracteoles with gland-fringed auricles. Flowers c. 12-15(7-20) mm
in diam.; buds cylindric, subacute. Sepals 5-7 x 0.7-1.7 mm,
subequal, narrowly lanceolate, aristate, with margin glandular-
ciliate and arista gland-tipped; veins 5, branched and reticulating
towards margin and distally; laminar glands pale, punctiform and
(mostly) black, shortly linear to punctiform, often dense; marginal
glands black, on short to long cilia. Petals bright? yellow, tinged or
veined red, 8-1 1 x 3-4- mm, c. 1.6 x sepals, oblong-lanceolate to
oblanceolate, rounded, without apiculus; laminar glands pale, punc-
tiform to shortly striiform, distal; marginal glands black, in dense
row, distal. Stamens (25?-)35-45, clearly 3-fascicled, longest 7-9
mm, c. 0.9 x petals; anther gland black. Ovary 3-locular, c. 2 x 1
mm, narrowly ovoid; styles 3, (3.5-)4-5 mm, 1.5-2.5 x ovary,
spreading-incurved or spreading. Capsule 3.5-4 xc. 2.5 mm, ovoid,
0.6-0.75 x sepals, enclosed by petals twisting together. Seeds
yellow-brown, c. 0.6 mm long; testa very finely scalariform (almost
smooth). 2n=18 (Reynaud, 1986), 16 (Love & Kjellqvist, 1974);
n=9 (Reynaud, 1986).

Streamsides, flushes and shaded places; 200-2000 m.

Spain (south-east: Andalucia, Murcia, Valencia, Castilla Nueva,
Aragon).

SPAIN. Granada: Sierra Nevada, Rio Jenil, 20 July 1 85 1 (fl), Bourgeau
1097 (K); Darro, 19 July 1883 (fl), Nilsson 1519 (BM). Jaen: Sierra de
Cazorla, Cueva del Peurco, 1070 m, 6 July 1951 (fl), Heywood 1040 (BM,
RNG). Murcia: Sierra Carrascoy, December 1855 (fl), Guirao in Bourgeau
s.n. (BM). Albacete: Sierra de Alcaraz, 2 km before turning to Riopar on
C415 from Alcaraz, c. 1 100 m, 7 August 1982 (fl), Goyder & Jury 338 (BM,
RNG); prope Alcaraz et Segura, 1000-2000 m, July 1890 (fl), Porta & Riga
II 400 (K). Valencia:/?^ Ramos (1993: 182). Castellon: Sierra de Segorbe,
351 m, August 1891 (fl), Reverchon s.n. (BM, FR). Cuenca: Solan, orillas del
Rio Cuervo, 14 August 1942 (fr), Cabalhro s.n. (K, MA*). Teruel: prope
Castelseras ad ripas aquosas fluminis Guadalupe, 30 July & 1 September

193

1975 (fl), Loscos Cent. I, 25 (FR). Burgos: Miranda de Ebro, orillas del Rio
Ebro, August 1929 (fl), Losa in Duffour 5855 (BM).

H. caprifolium is easily distinguished from the North African 14. H.
naudinianum by the aristate sepals. From the scanty available
evidence, the chromosome number is smaller (2n=16, not 18).

Subsect. 4. Adenosepalum (see p. 90). Type: H. montanum

Shrubs or perennial herbs with indumentum to base of inflores-
cence, or rarely stem or leaves or whole plant glabrous; leaves free;
bracts and bracteoles usually glandular-auriculate. Spp. 1 6-24.

16. Hypericum reflexum L.f., Suppl. pi: 346 (1781); Aiton, Hon.
kew. 3: 106 (1789), 2nd ed. 4: 179 (1812); Lam., Encycl. 4: 162
(1797); Willd., Sp. pi. 3: 1458 (1802); Buch in Abh. K. Wss.
Berlin 1816-1817: 362, 37 1 , 380 ( 1 8 1 7); Choisy, Prodr. monogr.
Hyperic.: 53 (1821), in DC., Prodr. 1: 551 (1824); Rchb.,
Iconogr. hot. exot. 1: 60, t. 86 ( 1 827); Buch, Phys. Beschr. Canar.
Ins. 153 (1828); Webb & Berth., Phytogr. canar. 1: 44 (1836);
Spach in Annls Sci. nat. (Bot.) II, 5: 357 ( 1 836); Masferrer mAn.
Soc. esp. Hist. nat. 9: 27 ( 1 880); Bornm. in Bot. Jahrb. 33: 453
( 1 903); Pitard & Proust, Les lies Canaries: 1 33 ( 1 909); R. Keller
in Engl. & Prantl, Nat. Pflanzenfam. 2nd. ed. 21: 179 (1925);
Stefanoff in God. Agr.-les. Fak. Univ. Sofiva 10: 58, t. 1 f. 5
(1932), 11: 149 (1933), 12: 83 (1934), in Pflanzenareale 4(1):
Karte 2b (1933); Ceballos & Ortuno, Veg. Fl.for. Canar. occid.:
387 (1951); Bramwell, D. & Z., Wildfls Canary Is: 70, f. 44
(1974); Kunkel, Endemismos Canarios: 296 (1977); Kunkel, G.
& M.A., Fl. Gran Canaria 4: 44, t. 165 (1979); N. Robson in
Cullen et al., Eur. Gdn Fl. 4: 72 (1995). Type: Canary Islands, no
loc., 1778 (fl), Masson s.n. (BM!-holotype). The younger
Linnaeus's 'Habitat in Barrancas Americae' must be regarded as
a lapsus calami.

Fig. 28B, Map 37.

H.foliosum sensu Brouss. ex Webb & Berth., Phytogr. canar. 1: 45
(1836) in synon.

H. reflexum var. leiocladum Bornm. in Bot. Jb. 33: 453 (1903)
['leioclada ']; Pitard & Proust, Les lies Canaries: 133 (1908); Lid
in Skr. norske Vidensk.-Akad. I, Math.-Nat. Kl., N.S. no. 23: 122
(1967). Types: Canary Islands, Gran Canaria, Tafira, Beo
Guiniguada, c. 400 m, n.d., Bornmiiller2l53 (JE-syntype); Gran
Canaria, Caldera de Bandama, 17 May 1900, Bornmuller 352
(JE-syntype).

H. reflexum var. myrtillifolium Bornm. in Bot. Jb. 33: 453 (1903)
['myrtillifolia']; Pitard & Proust, Les lies Canaries: 133 (1908)

Map 37 Sect. 27: 13. H. coadunatum A (see also Map 36); 16. H.
reflexum (limits on Tenerife according to Voggenreiter, 1974) specimens
•, record O.

194

['myrtifolia']. Type: Canary Islands, Teneriffe, ad mare prope
Taganana, n.d., Bornmuller s.n. (JE-holotype).
H. reflexum var. lanuginosum Pitard in Pitard & Proust, Les lies
Canaries: 133 (1908) ['lanuginosa'], in Fedde, Repert. Spec.
Nov. 9: 210 (191 1); Lid in Skr. norske Vidensk.-Akad. I, Math.-
Nat. Kl., N.S. no. 23: 122, f. 9b (1967). Type: Canary Islands, 6
specimens cited; Tenerife, Guimar Barranco de Badajoz, 400 m,
n.d., Pitard s.n. (TUB-lectotype, selected here).

Icones: Rchb., Iconogr. hot. exot. 1: t. 86(1827); Bramwell, D. &Z.,
Wild/Is Canary Is: f. 44 (1974); Kunkel, G. & M.A., Fl. Gran
Canaria4: t. 165(1979).

Shrub 0.5-1 m tall, densely and divaricately branched below, ±
bushy, with branches erect to ascending, the whole plant (usually
except stems) glabrous. Stems red-brown, compressed (sometimes
shallowly 2-lined) and densely villous-tomentose or rarely sparsely
puberulous to glabrous when young, soon terete, eventually gla-
brous; internodes shorter than leaves; cortex exfoliating in usually
internodal sheets; bark dark red-brown, smooth. Leaves sessile;
lamina 10-25 x 3-12 mm, narrowly elliptic or narrowly oblong to ±
broadly triangular-lanceolate, concolorous or slightly paler beneath,
chartaceous, not or slightly glaucous, glabrous, plane, spreading or
somewhat deflexed; apex acute to subacute, margin entire, base
cordate to rarely rounded and ± deeply amplexicaul; venation: 2(3)
pairs of laterals curved-ascending from lower 0.25(-0. 35) of midrib,
tertiary reticulation dense and manifest but not prominent; laminar
glands pale, dense to rather sparse, unequal, ± prominent;
intramarginal glands all black or a few pale, dense but sometimes
irregular. Inflorescence c. 5^K)-flowered from up to 3 nodes, with-
out flowering branches from lower nodes, curved corymbiform and
dense or rarely lax and rounded-pyramidal; pedicels 2.5^ mm;
bracteoles (and bracts at uppermost 2-3 nodes) narrowly triangular-
lanceolate to linear, with rather sparse black-glandular cilia. Flowers
c. 15-20 mm in diam.; buds narrowly ellipsoid, acute. Sepals 3.5-6
x 1 -2 mm, subequal, free or to c. 0. 1 5 united, lanceolate to narrowly
oblong, acute to subacuminate, with sessile marginal glands or
margin glandular-ciliate; veins 3, laterals branching; laminar glands
pale, striiform to usually punctiform; marginal and inframarginal
glands black, round- or flat-topped, immersed to sessile or on short
(rarely long) cilia. Petals rather pale bright yellow, not tinged red,
10-12(-14) x 4-5 mm, c. 2.5 x sepals, oblong-oblanceolate, ob-
liquely truncate to rounded, apiculus obsolete or absent; laminar
glands pale, few, linear (rarely) or striiform to punctiform; marginal
and inframarginal glands all black or some pale, not prominent.
Stamens c. 25, longest 9-10 mm, c. 0.8-0.9 x petals; anther gland
amber. Ovary 2-2.5 x 1-1 .5 mm, narrowly ovoid; styles 6-7 mm, c.

N.K.B. ROBSON

3 x ovary, widely spreading-incurved. Capsule 4-5 x 2.5-3.5 mm,
ovoid-ellipsoid, shorter than or equalling sepals, enclosed by petals
twisting together. Seeds yellow-brown, c. 0.6 mm long; testa
shallowly linear-foveolate. 2n= 18 (Larsen, 1962;Borgen, 1969; van
Loon & de Jong, 1978, as var. lanuginosum; Reynaud, 1986).

Damp cliffs and walls, around springs and along rocky streams in
open localities; 150-1600 m.

Canary Islands (all western islands, i.e. not Fuerteventura or
Lanzarote).

CANARY ISLANDS. Tenerife: in rupibus convallis Bufodero, 1 6 April
1 855 (fl), Bourgeau 1 239 (K); in vicinitate pagum Santa Ursula, Barranco de
la Crux, c. 200 m, 1 8 July 1 933 (fl), Asplund 855 (BM, K); Monte Verde, c.
20 km along Orotava - Canada road, 1300 m, 29 August 1971 (fl), Chicken
8 1 (BM). Gomera: Barranco de la Laja, 28 May 1 894 (fl), Murray s.n. (K);
near Benchijiqua, 600 m, 9 May 1977 (fl), Jarvis 638 (BM). La Palma: no
specimens of H. reflexum s.s. seen, only two of H. xjoerstadii (q.v.). Gran
Canada: prope Tafira, 400-500 m, 1 6 May 1 900 (fl & e. fr), Bornmuller 35 1
(H); Risco Blanco de Tirajana, 14 May 1975 (fl), Bramwell & Humphries
JC240(BM);ValledeAgache,SWside, 1000m, 1 August I960 (fl), Andrews
R40 (K).

H. reflexum is apparently not closely related to 1 . H. glandulosum,
although the occurrence of intermediates (Ix. H. x joerstadii)
suggests that they hybridize. The nature of these intermediates might
be revealed by cytological investigation, as the species can have
different chromosome numbers, viz. H. glandulosum 2n=40, 1 8, H.
reflexum 2n=18.

H. reflexum is variable, but there are no breaks in variation such as
would warrant the recognition of plants with glabrous stems (var.
leiocladum Bornm.)12 or short, ovate or ovate-elliptic leaves (var.
myrtillifolium Bornm.) as distinct taxa.

H. reflexum is the basic species to which the H. montanum group
(Spp. 17-24) is related. The chromosome numbers so far recorded
for species in this group are all 2n=16 or 32.

17. Hypericum montanum L., Fl. Suec. 2nd ed.: 266 (1755), Sp.
pi. 2nd ed.: 1 105 (1762); Oeder, Fl. Danica 1: t. 173 (1764);
Smith, Engl. Bot. : t. 37 1 ( 1 797?); Ucria, Hon. reg. panorm. : 326
(1809); Choisy, Prodr. monogr. Hyperic.: 54 (1821), in DC.,
Prodr. 1: 55 1 ( 1 824); Spach, Hist. nat. veg. Phan. 5: 392 ( 1 836),
in Annls Sci. nat. (Bot.) II, 5: 357 (1836); Rchb., Ic. fl. germ,
helv. 6: t. 347 f. 5187 (1844); Syme, Engl. Bot. 3rd ed.: 158, t.
275 (1863); Boiss., Fl. orient. 1: 807 (1867); Willk. & Lange,
Prodr. fl. hispan. 3: 593 (1878); Batt. & Trab., FL Algerie,
Dicots. 1: 182 (1888); Maire in Mem. Soc. Sci. nat. Maroc, no.
7 (1924); Hayek, Prodr. fl. pen. bale. 1: 539 (1925); R. Keller in
Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 1 19 (1925); Hegi,
///. Fl. Mittel.-Europa 5(1): 524, t. 183 f. 3, ff. 2009, 2010
(1925); Jahand. & Maire, Cat. Pis Maroc 2: 482 (1932); Stefanoff
in God. Agr.-les. Fak. Univ. Sofiya 10: 58, t. 3 f. 36 (1932), 11:
165 (1933), 12: 85 (1934); Gorshk. in Shishkin & Bobrov, Fl.
U.R.S.S. 15: 245 (1949); Guinea, Viscaya Paisaje veg.: 219 &
map (1949); Ross-Craig, Drawings Br. Pis 6: t. 15 (1952);
Grossh., Fl. Kavk. 2nd ed. 6: 174, Karta 192 (1962); N. Robson
in Davis, Fl. Turkey 2: 385 (1967), inTutin et al., Fl. europaea 2:
265 (1968); Hulten, Atlas vdxt. utbred. Norden: 317, map 1231
(197 1 ); Kask inEestiN.S. V. Floora 8: 28, ff. 6-8 ( 197 1 ); Franco,
Nova Fl. Portugal 1: 450 (1971); Stjep.-Vesel. in Josifovid, Fl.
Srbije 3: 112, t. 32 f. 3 (1972); Fournier, Quatre flores France
2nd ed.: 455 (1977, repr. 1990); Meusel, Vergl. Chor. Zentraleur.
Fl. Texte 2: 23, Karten 2: 284 (1978); Mennema,

12 But seep. 175.

STUDIES IN HYPER1CUM

195

Fig. 28 A. H. montanum: (a) habit; (b) leaf (lower surface) and part of stem; (c) bract; (d) sepal; (e) petal; (0 capsule. B. H. reflexum: (g) habit; (h) leaf
(lower surface) and part of stem; (i) sepal; (j) petal; (k) capsule (a, b, g x Vr, h x 2; c-f, i-k x 3). A. Font Quer her maroc. 1928 268. Lacaita 6392. B.
Bourgeau 80, Asplund 855.

196

N.K.B. ROBSON

Quene-Brot. & Plate, Atlas Netherl. Fl. 1: 132 (1980); Pignatti,
Ft. Italia 1: 346 (1982); I. Hagemann in Flora 173: 1 17, ff. 19,
39 (1983); Ramos in Trab. Dep. Bot. Univ. Complut. 12: 54
(1983), inActabot. Malacit. 11: 169, f. 9c (1986); N. Robson in
Wild Fl. Mag. no. 403: 17 (1985); Lid, Norsk-Svensk-Finsk FL:
3 1 6, 74 1 ( 1 985); Greuter, Burdet & Long, Med-Checklist 3: 269
(1986); Clapham, Tutin & Moore, Fl. Br. Isles 3rd ed.: 116
( 1 987); Haeupler & Schonf., Atlas Farn. -u. Bliitenpfl. Bundesrep.
Deutschl.: 330, Karte92(1988); Perring & Walters, Atlas Br. FL,
repr. amend.: 59 ( 1 990); Stace, New FL Br. Isles: 256 ( 1 99 1 ); N.
Robson in Cullen et al., Eur. Gdn FL 4: 72 (1995). Type:
Sweden, Skane?, 'in montibus Westrogothiae 213. Scaniae',
Herb. Linn. 943.39 (LINN'.-lectotype, selected here;
SBT[labelled by Linnaeus, fide Jarvis]-syntype). Although
Linnaeus's protologue includes references to Bauhin's Pinax
and Historia, Columna's Ekphrasis and Fuch's Historia, the
absence of this plant from the first edition of Species plantarum
(1753) suggests that either Linnaeus had not seen material of it
by that date or (more likely) that he had overlooked it. From the
relatively full treatment of//, montanum in both FL suecica 2nd
ed. and Species plantarum 2nd ed., however, it would seem that
by 1755 he had obtained such material. The only specimen of
this species in the Linnaean Herbarium (LINN) is unannotated,
apart from the name; but there seems little doubt that it is one of
the specimens mentioned in Fl. suecica. It is therefore the most
appropriate lectotype.
Fig. 28A, Map 38.

H. elegantissimum Cr., Stirp. austr. 2: 63 (1763). Type: Austria, 'In
Gatterholzel, Dornbach & alibi non infrequens', Crantz (BP!-
holotype).

H. glandulosum Gilib., Fl. Lituan. 2: 205 (1782), nom. illegit.
superfl. (based on H. montanum L.).

H. confertum Moench, Meth. hot.: 129 (1794), nom. illegit. superfl.
(based on H. montanum L.).

H. montanum [var.] (3 triphyllum Choisy, Prodr. monogr. Hyperic.:
54 (1821), in DC., Prodr. 1: 552 (1824). Type: France,
Fontainebleau, Anon. (G-DC!-holotype).

H. montanum [var.] (3 scabrum Koch, Syn. fl. germ. helv. 1: 135
(1835); Hayek, Prodr. fl. pen. Bale. 1: 539 (1925); Hegi, ///. FL
Mittel-Europa 5(1): 525 (1925); Maire in Bull. Soc. Hist. nat.Afr.
Nord 27: 216 (1936) et auct. plur. Type not stated.

H. montanum [var.] p sensu Ledeb., Fl. rossica 1: 450 (1842) = var.
caucasicum Boiss. (see below).

H. tauricum sensu hort. ex Ledeb., Fl. rossica 1: 450 ( 1 842) in synon.

H. montanum [var.] (3 caucasicum Boiss., FL orient. 1: 807 (1867);
Parl., FL Hal. 5: 532 (1875); Woronow in Kuzn., Busch & Fomin,
FL Cauc. crit. Ill, 9: 46 (1906); Hegi, ///. FL Mittel-Europa 5(1):
526 (1925) ['var. caucasicum Parl.']; Grossh., FL Kavk. 3: 70
(1932), 2nd ed. 6: 174 (1962). Types: Turkey, Trabzon, vallee
d'Of (Lazistan), vers 200 m, 27 June 1866 (fl & fr), Balansa 86
(G! -lectotype; E!-photograph); Georgia, Guria, n.d. (fl), Szowits
s.n. (LE-syntype; G!, E! -photograph).

Adenosepalum montanum (L.) Fourr. in Ann. Soc. Linn. Lyon N.S.,
16:353(1868).

Map 38 Sect. 27: 17. H. montanum specimens •. records O; limits according to literature.

STUDIES IN HYPERICUM

197

Hypericum montanum forma humifusoides Kuntze in Flora 1880:

305 ( 1 880). Type: Germany, Saxony, Spitzberg bei Wurzen, 1 879,

Kuntze (NY-holotype).
H. montanum [var.] a typicum G. Beck, Fl. Nieder-Osterr.: 531

(1892); Hegi, ///. Fl. Mittel-Europa 5(1): 525 (1925). Type as for

H. montanum L.
H. montanum var. scaberulum G. Beck, Fl. Nieder-Osterr.: 531

( 1 892), nom. illegit. superfl. (based on//, montanum var. scabrum

Koch).
H. ciliatum var. pseu dociliatum R. Keller inAlbov, Prodr.fl. colchic.:

42 (1895). Type: Georgia, Guria, jugum Adzharo-Imereticum,

1893, Ardasenov s.n. (G?-holotype).
H. montanum forma ternatum Borbas, A Balaton Tudom. Tunul.

Ered. 2. Szakasz: 403 (1900). Type not indicated.
H. perfoliatum var. pseudociliatum (R. Keller) Woronow in Kuzn.,

Busch & Fomin, Fl. Cauc. crit. Ill, 9: 49 (1906).
H. montanum forma abbreviatum Reinecke in Mitt. Thiir. Bot. Ver.

30: 19(1913), in Jb. K. Akad. gem. Wiss. Erfurt 40: 150 (1914).

Type: Germany, ThUringen, Erfurt, Mobisburger Holze siidlich

von Rhoda, Reinecke (JE?-holotype).
H. montanum subsp.? elegantissimum (Cr.) G. Jav., Mag. Fl. 2: 720

(1925).
H. montanum forma subprolificum Murr (\92%),fide Soo (1968).

Icones: Syme, Engl. Bot. 3rd ed.: t. 275 (1863); Ross-Craig, Draw-
ings Br. Pls6:t. 15(1952).

Perennial herb 0.2-0.8 m tall, wholly erect or decumbent to ascend-
ing but not rooting at base, with woody taproot, few-stemmed,
usually unbranched below inflorescence, wholly glabrous often
except leaves beneath. Stems green to reddish, terete; internodes all
or upper exceeding leaves. Leaves sessile; lamina (20-)25-70 x
(10-)13-28 mm, oblong-elliptic to lanceolate or ± broadly ovate,
paler beneath, thinly chartaceous, not glaucous, wholly glabrous or
usually scabrid to puberulous beneath, plane, spreading; apex ob-
tuse to rounded or uppermost subacute, margin entire, base rounded
to truncate or subcordate; venation: 3^t pairs of laterals curved-
ascending from lower 0.2-0.25 of midrib; tertiary reticulation dense,
not or slightly prominent; laminar glands pale, dense, unequal, not
prominent, rarely absent ('var. caucasicum'); intramarginal glands
black, dense but irregular, the larger often submarginal. Inflores-
cence 7-32(-c. 50)-flowered from up to 3(4) nodes, without flowering

branches from lower nodes, curved-pyramidal or corymbiform to
subcapitate and dense, or lower node(s) ± distant with dense partial
inflorescences, the whole then narrowly pyramidal to narrowly
cylindric; pedicels 1.5-4 mm; bracts and bracteoles linear, black-
glandular-ciliate, densely glandular-auriculate. Flowers 10-15(-20)
mm in diam.; buds narrowly ellipsoid to cylindric, rounded. Sepals
5-6 x 1-1.5 mm, subequal or equal, free or very shortly united,
narrowly oblong to narrowly lanceolate, acute, with margin rather
long-glandular-ciliate; veins 5, laterals branching; laminar glands
pale, linear to striiform; marginal glands black, flat-topped. Petals
pale yellow, not tinged red, (8-) 10- 12 x 2-3.5 mm, c. 2 x sepals,
oblong-lanceolate, rounded, apiculus absent; laminar glands pale,
few, elongate-punctiform or usually absent; marginal glands absent.
Stamens 20-28, longest 6-9 mm, c. 0.75 x petals; anther gland
black. Ovary 2.5-3.5 x 1.5-2.5 mm, narrowly to rather broadly
ovoid; styles 3-4 mm, 1.2-1.35 x ovary, spreading. Capsule 6-8 x
4-6 mm, ovoid, exceeding sepals, enclosed when developing by
petals twisting together. Seeds dark red-brown, 0.8 mm long; testa
linear-reticulate. 2n=16 (Noack, 1939; Robson, 1956; Reynaud,
1975; Love & Love, 1982), n = 8 (Nielsen, 1924; Gagnieu &
Wilhelm, 1965).

Woods, thickets and hedgebanks, on calcareous or gravelly soils;
lowland (especially in the north) to 1200 m (Italy), 1450 m (Spain)
and 1950 m (Morocco).

From northern England, Denmark, southern Norway, and southern
Sweden, southern Finland (one locality), Estonia (one locality),
Poland, Belarus and the Ukraine (including Krym), south to N.
Portugal, central Spain, Corsica, Sardinia, Italy, Bosnia, Serbia and
Romania; also Georgia, NE Turkey and Morocco.

GREAT BRITAIN. Devon: Brixham, Berry Head, 19 July 1936 (fl),
Hall s.n. (BM). Surrey: Addington, Court Wood, 31 July 1924 (fl), Salmon
s.n. (BM). Oxfordshire: near Caversham, 3 August 1890(fl), Wolley Dod s.n.
(BM). Gloucestershire (W.): Tidenham, August 1922 (e. fr), Redgrave s.n.
(BM). Glamorgan: Bishopston Valley, August 1913 (fr), Shepherd s.n. (BM).
Pembroke: Tenby, Black Rock, 19 August 1954 (fr), Townsend s.n. (K).
Caernarvon: Great Orme's Head, 18 July 1912 (fl), Bickham s.n. (BM).
Yorkshire (NE): Helmsley, 2 August 1952 (fl & fr), Bangerter 125 (BM).
Westmorland: Anna Well Force, 5 August 1927 (fl), Foggitt s.n. (BM).

NORWAY. Hordaland: Bergen, Hardanger, Godoii, 8 August 1 982 (fl &
fr), Rettig s.n. (H). Vest Agder: Randesund Dvergones, 9 August 1959 (e. fr),
Skrimer s.n. (H).

SWEDEN. Goteborg: Partille, L. Prasttjarn, 30 July 1928 (fl), Ohlsen in
Samuelsson PI. Suec. 1 155 (BM, K). Alvsborg: Njorn, Sundsby, 9 July 1950
(fl), Hayren-Malmstrom s.n. (H). Jonkoping: in montibus Huskvarna, Sep-
tember 1902 (fr), Bergstrand s.n. (H). Ostergotland: Valdemarsvik, Borg, 5
August 1985 (1. f\),Nannfeldt 19047 (BM). Halland: Halmstad, 27 July 1956
(fl), P. Fries s.n. (H). Malmohus: Helsingborg, 26 July 1885 (fl), Trolander
(H). Gotland: Slite Tildehajdal, 20 July 1968 (fr), Alanko (H).

FINLAND. Turku: Abo, Lojo, juxta praed. Luusi, in pago Jantoniensi,
1 1 August 1915 (fl), Lindberg in PI. Fin. Exsicc. 1241 (K).

ESTONIA. Recorded from Saaremaa I. (Osel), Harva (Kask, 1971: 30,
f.7).

DENMARK. Jylland: Ribe Varde, 26 August 1896 (fr), Boldt s.n. (H);
Viborg,Daugbjerg,W. ofViborg (distr. 15),4August 1974 (fl), ATJM/Z s.n. (H).
Sjaelland: Nordsjaelland, Holte, Geelskov, 3 July 1943 (fl), Dahl D. 45a
(BM). Mon: Freuchens Pynt, 3 July 1935 (fl), von Wendt s.n. (H).

GERMANY. Niedersachsen: Hildesheim, July 1867 (f\),Evers501 (H).
Nordhein-Westfalen: Bonn, n.d. (fl & fr), Sievreck s.n. (H). Rheinland-Pfalz:
Donnersberg, 28 July 1905 (fl), Knabes.n. (H). Hessen: UnteresWerragebiet,
29 July 1 898 (1. fl), Goldschmidt s.n. (FR). Baden-Wurtemberg: Tubingen, 19
July 1922 (fl), Petterssen s.n. (H). Bayern: Steinebach am Worthsee, 1
September 1918 (fr), Oberneder32&3 (BM).Thiiringen: bei den Dornburger
Schlossern, 9 July 1950 (fl), Launert s.n. (BM). Sachsen-Anhalt: Alte
Stolberg, 8 July 1882 (fl), Vock s.n. (K). Mecklenburg: Crivitz, 12 July 1880
(fl), Neverrmann s.n. (BM).

198

N.K.B. ROBSON

HOLLAND. Scattered records in south and south-east (fide Mennema,
Quene-Broterenbrood & Plate, 1980: 132).

BELGIUM. Namur: Nettine, July 1937 (fl), Masseray s.n. (BM).
Hainault: Obourg, 4 July 1867 (fl), 22 August 1867 (fr), Martinis VI 260
(BM, K).

LUXEMBURG. Several records (fide Rompaey & Delvosalle, 1979:
map 423).

FRANCE. Seine-et-Oise: Coteau de la Ferte-Alais, 23 July 1876 (fr),
Bwiw/s.n.(K).Loir-et-Cher:CoteaudeLavardin, 1856(fr), Vore/631 (BM).
Maine-et-Loire: Soucelles, 16July 1874(1. fl), Geneviers.n. (BM). Morbihan:
Belle lie vis-a-vis Kerouarh, Pont-en-Dro, 3 July 1901 (fr), Gadeceau 631
(BM) -extinct? Deux-Sevres: foret de Chize, 24 July 1910(fr), Gadeceau s.n.
(BM). Cote d'Or: Messigny, Combe de Chainoux, 1 8 July 1 869 (fr), Laguesse
s.n. (K). Haut-Rhin: Habsheim, foret de la Hardt, 240 m, 20 July 1974 (fr),
Rastetter in Exsicc. Auguier F. 16 7616 (BM). Doubs: Bonnevaux, n.d. (1. fl),
Depierre s.n. (BM). Ain: Miribel, July 1912 (fl), Reverchon s.n. (FR). Haute-
Savoie: Vacheresse, a la Baume, 30 July (fr), Depierre s.n. (BM). Savoie:
Mont Grelle pres Chambery, 9 August 1896 (fr), Borel s.n. (K). Basses-
Alpes: Goudeissart pres Barcelonnette, 1 1 August 1 884 (fl), Wilmott s.n. (K).
Var: Mont des Maurs, near Collobrieres, 24 June 1914 (fl), Adamson s.n.
(BM). Isere: Villard-de-Lans, Gorge de la Bourne, 30 September 1949 (fr),
Sandwith 3529 (K). Rhone: Lyon a Niron, 3 July 1 883 (fl), Jordan s.n. (BM).
Haute-Loire: de Lempdes a Blesle, June 1876 (fl), Girardet s.n. (BM).
Cantal: environs d'Ydes, Lisiere des bois, 18 August 1873 (fr), A. Braun s.n.
(BM). Aveyron: Viviez, June 1905 (fl), Garrau in Elias%\2 (BM). Pyrenees-
Orientates: Vernet, 900 m, 9 July 1934 (e. fr), Wyatt 1 19 (K). Ariege: Ax les
Thermes, July 1925? (fl), Lofthouse s.n. (BM). Haute-Garonne: Luchon, 14
July 1 889 (fl), Murray s.n. (BM). Hautes-Pyrenees: zwischen St Sauveur und
Sazos, 3 July 1925 (fl), Ronniger s.n. (W).

SPAIN. Oviedo: Oviedo, gorge of R. Sella, 13 July 1927 (fr), Wilmott
s.n. (BM). Santander: Picos de Europa, supra pagum Epinama, c. 1000 m,
September 1930 (fr), Buck s.n. (H). Huesca: Los Aranones (Caufranc), 2
August 1955 (fl & fr), Sandwith 4488 (K). Lerida: Val d'Aran, Foret de
Betren, 1200 m, 25 July 1934 (fr), Estiva! in Sennen 9056 (BM). Gerona: Sol
de Santicosa, 950 m, 8 August 1934 (fr), Sennen s.n. (BM). Barcelona: in
dumetis Montiserrati, July 1906 (fr), Marcet s.n. (BM). Madrid: Kast.
Schiedegebirge [Sade Guadarama], Somosierra, 'Hayedo de Montejo', 1 350
m, 6 June 1962 (st), Em s.n. (FR).

PORTUGAL. Tras os Monies: Braganc.a, Monte de San Bartolomeu, 24
June 1955 (fl), Fernandes, Matos & Matos 5488 (BM).

CORSICA. Ajaccio to Curti [Corte]: Vivazione, 500-700 m, 3 July 1 970
(fl), Verdcourt41%\ (K); ?: Premier Cascade del' Aqua ardente, 13July 1917
(fl), Forsyth Major H292-23 (K).

SARDINIA. No specimens seen, but recorded by e.g. Pignatti (1982).

ITALY. Venezia: Resiutta, slopes of Plauris, 17 July 1863 (1. fl), Hb.
Churchill s.n. (K).Trentino/AltoAdige: Molveno, 4August 1926 (fr), Barton
s.n. (BM). Lombardia: Chiavennas, 9 July 1886 (fr), Murray s.n. (BM). Val
d'Aosta: Val de Cogne, over Valoutay, 3 August 1864 (fr), Hort s.n. (BM).
Piedmonte: pratis alpinis Valdensium, July 1860 (fr), Rostan s.n. (BM).
Liguria: Portofino Peninsula, S. of Genoa, 10 September 1963 (1. fl), Robson
1807 (BM). Emilia-Romagna: Modena, Piandelagotti, Boschi, 1150 m, 4
July 1934 (fl), Lennart s.n. (H).Toscana: Gugena, 29 June 1964 (fl), Ranhala
s.n. (H). Abruzzi e Molise: M. Sirente, 1 500 m, July 1 875 (fl), Groves s.n. (K).
Campania: Salerno, Monte Mai de Calvanico, c. 1200 m, 19 July 1921 (fl),
Lacaita 240/21 (BM).

SWITZERLAND. Geneve: prope Genevam, n.d. (fr), Herb. Boissier
(K). Vaud: La Comballaz, 10 July 1881 (fl), Branner s.n. (H). Valais: Val
d'Entremont, Sembranchori, 880 m, 16 July 1980 (fl), Lawalree 22569
(BM). Bern: Meiringen, Starelamm, Finstere Schucht, 4 September 1924 (fl
& fr), Barton s.n. (BM). Zurich: Sihtwald, 21 July 1923 (fl), Hayren s.n. (H).
Schwyz: near Brunnen, 27 July 1898 (fl), LS. Wright s.n. (K). Glarus:
Klontal, 21 August 1930 (fl), Lacaita 6391 (BM). Ticino: Monte Generoso,
6 July 1856 (fl), Murray s.n. (BM).

AUSTRIA. Tirol: Lienz, 20 July 1869 (fl), Gander s.n. (BM, K).
Steiermark: Graz, Racherkogel, c. 500 m, July 1 9 1 1 (fr), Fritsch in Hayek Fl.
Stir, exsicc. 1201 (BM, H). Niederosterreich: Semmering, Reichenau, Au-
gust 1924 (fr), Buch s.n. (H).

HUNGARY. Several localities (fide Soo, 1968: 437).

SLOVENIJA. Postojna, 300 m, 20 July 1 960 (e. fr), McCallum Webster
4081 (K).

CROATIA. Istra, Abbazia, oberhalb Vrutki-Tal, 25 June 1935 (fl),
Ronniger s.n. (W).

SERBIA and BOSNIA HERZEGOVINA. Present (fide Stepanovi6
Vesilichi£, 1972).

ROMANIA. Transylvania: Arad, p.p. Camna, 8 September 1974 (fr),
Danciu s.n. (BM); Covasna, prope opp. Sf. Gheorghe, 14 August 1981 (fr),
Danciu s.n. (BM); Brasov, p.p. Cristian, c. 800 m, 1 August 1973 (fr),
Parascan & Danciu s.n. (BM).

CZECH REPUBLIC. Moravia: M. Weisskirchen, Parsihouster Reviers,
August 1912 (fr), Petrak Fl. Boh. & Mor. XI 1048 (BM); Briinn [Brno],
Runitz, July 193 1 (fl), Hruby s.n. (K). Also in Bohemia and Slovakia.

POLAND. Poznan: Konen, 16 July 1868 (fr), Baenitz s.n. (BM).

BELORUSS YA. Brest, bei Sevcki (?), 20 June 1 859 (fl), Lehmann s.n.
(BM).

UKRAINE. Lesienice k. Lwowa [Lvov], 19 July 1938 (fr), Madalski PI.
Pol. Exsicc. 332 (BM, K); Bojarka, 1 July 1903 (fl), 15 July 1903 (fr), Finn
169 (H). Krym: Kastali (Vulf, 1953).

GEORGIA. Poti, n.d. (fl & fr), Nordmann 308 (H); Guria, n.d. (fl),
Szowitz (G, E-photograph).

TURKEY. Giresun: 42 km S. of Giresun (N. of Khumbet), 1000 m, 7
July 1969 (fr), Sorger 69-24-4 (W).Trabzon: Lazistan, Vallee d'Of, vers 200
m, 27 June 1 866 (fl & fr), Balansa 86 (G, E-photograph); Sumela Kloster,
800-1 300 m, 28 July 1 982 (fr), Sorger & Buchner 82-9 1-1 8 (W). Rize: ? (no
specimens seen). Coruh: Artvin and Ardanu? (Grossheim, 1962: map 192).

MOROCCO. Rif: Mt Buhaschen [Yebel Buhasen], 1300 m, 20 June
1928 (fl), Font Quer Iter Maroc. 1928 268 (BM); Ketama, Telata, vers 1450
m, 5 August 1932 (fl), Sennen & Mauricio s.n. (BM). Za'i'an: Azrou, 12 km
from Azrou to Ifrane, 1350 m, 26 June 1974 (fl), Reading U./B.M. Exped.
1058 (BM,RD*).Moyen Atlas: Foret d' Am Kahla, 1950m, 20 July 1924 (fr),
Jahandiez 832 (BM).

H. montanum is the north-western member of a pair of species that
appears to be directly related to 16. H. reflexum (from the Canary
Islands), the other member being 1 8. H. annulatum. H. montanum is
mainly west, central and east European and north-west African; H.
annulatum is mainly south-east European and north-east to east
African.

H. montanum can be separated from H. annulatum only on a
combination of characters, since the latter in particular is very
variable. H. montanum is always glabrous except on the lower leaf-
surface, which is usually scabrid. Where that, too, is glabrous (var.
typicum G. Beck), the species can nearly always be distinguished
from wholly glabrous forms of H. annulatum by the condensed
inflorescence or partial inflorescences. In most cases, however, H.
annulatum can be recognized by the presence of hairs on the stem
and upper leaf-surface and the relatively lax inflorescence.

H. montanum is not very variable other than in leaf shape, the
indumentum of the lower leaf-surface and the presence or absence of
partial inflorescences. The scabrid lower surface of the leaf (var.
scabrum Koch, var. scaberulum G. Beck) is by far the commoner
state, although Beck named the wholly glabrous form var. typicum.
The lectotype, too, is scabrid beneath, and so var. typicum is not the
type variety, i.e. var. montanum. In fact, the glabrous leaf seems to be
no more than a shade-induced state that does not merit taxonomic
recognition.

In Georgia and Turkey most of the populations have leaves
without laminar pale glands (var. caucasicum Boiss.); but this
character is not wholly constant, and so I have not given these
populations taxonomic recognition either.

Finally, H. montanum var. pilosum Horwood is H. hirsutum L.,
whilst var. maculantherum Sagorsky and var. punctatum Andreanszky
are almost certainly respectively H. spruneri Boiss. and H.
perfoliatum L., although I have not yet managed to see the type of
either variety. The record of H. montanum from southern Greece
(Messinia) (Sibthorp & Smith, Fl. Graecae Prodr. 2: 117, 1813)
possibly refers to H. vesiculosum Griseb. (sect. 13. Drosocarpium),
which occurs there.

STUDIES IN HYPERICUM

199

18. Hypericum annulatum Moris, Stirp. sard, clench.: 9 (1827);
Fl. Sardoa 1: 323, t. 22 (1837); Walp., Repert. hot. syst. 1: 384
(1840); N. Robson in Kew Bull. 12: 444 (1958), in Fl. Europaea
2: 265 (1968); Moggi & Pisacchi in Webbia 22: 212, f. 13
(1967);Arrig. et al. in Webbia 28: 423 & f. 1 (1973); Pignatti, Fl.
Italia 1: 46 (1982); Chiappini, Fl. paesag. veg. Sardegna: 26, t.
11 (1985); Greuter, Burdet & Long, Med-Checklist 3: 264
(1986); N. Robson in Cullen et al., Eur. Gdn Fl. 4: 72, ff. 1 1 .5,
11.15 (1995) (= '//. degenW). Type: Sardinia, in summo S.
Vittoria esterzili, July 1826 (fl), Moris s.n. (TO-holotype, K!-
photograph; Fl- isotype).

Fig. 29C (subsp. afromontanum).

Perennial herb 0.2-0.75 m tall, wholly erect or shortly decumbent
but not rooting at base, with ± woody taproot, usually few-stemmed,
unbranched below inflorescence or sometimes with short axillary
shoots, wholly glabrous or usually puberulous to densely shortly
whitish-pubescent on stems below inflorescence and on both leaf
surfaces. Stems green to reddish, terete; internodes all shorter than
leaves or upper exceeding them, eglandular or sparsely (along
'lines') to densely black-gland-dotted or shortly black-gland-
streaked. Leaves sessile; lamina (5-)10-55 x (3-)5-32 mm, narrowly
oblong or oblong-elliptic or lanceolate or rather broadly ovate, paler
beneath, thinly chartaceous, not glaucous, glabrous to ± densely
pubescent on both sides, plane, spreading; apex acute to rounded,
margin entire, base rounded to truncate or subcordate; venation:
(3)4-5 pairs of laterals curved-ascending from lower 0.2-0.5 of
midrib, tertiary reticulation dense, not prominent; laminar glands all
pale or occasionally some black, dense, unequal; intramarginal
glands black, dense but irregular to rather sparse. Inflorescence 5-c.
120-flowered from up to 4 nodes, without or very rarely with 1-2
flowering branches from next lower node, pyramidal or shortly
cylindric to corymbiform, lower nodes not distinct, the partial
inflorescences nearly always lax-flowered when mature; pedicels 1-
4 mm; bracts and bracteoles linear, black-glandular-ciliate, densely
glandular-auriculate. Flowers 15-25 mm in diam.; buds cylindric-
ellipsoid, obtuse. Sepals (4-)5-6(-8) x 1-1 .5 mm, subequal, free or
almost so, narrowly oblong to narrowly lanceolate, acute or very

rarely subacute, with margin long- to short-glandular-ciliate; veins 5
or 3 with strong near-basal outer branch from each lateral, these
sometimes united (commissural) at base; laminar glands all pale to
all black, linear at base or all striiform to punctiform; marginal
glands black, flat-topped. Petals pale (?) to golden yellow, some-
times veined or tinged red, (8-)10-13(-15) x (2.5-)3-3.5 mm, c. 2
x sepals, oblong-lanceolate, rounded, apiculus absent; laminar glands
pale and/or black, scattered; marginal glands absent. Stamens c. 20-
40, longest (7.5-)9-10 mm, 0.65-0.8 x petals; anther gland black.
Ovary 2-3 x 1-2 mm, narrowly ovoid-pyramidal; styles 6-8 mm, 2-
4 x ovary, spreading-incurved. Capsule (4-)5-8 x 2.5-4 mm, ovoid,
equalling or exceeding sepals, enclosed when developing by petals
twisting together. Seeds dark yellow-brown, 0.6-0.7 mm long; testa
linear-reticulate. 2n = 16 (Reynaud, 1980; Strid & Franzen, 1981:
836).

Dry or stony places, in scrub or grassland; 1212m (Sardinia), 300-
c. 1330 m (Balkans), 1050-1725 m (Arabia), 1600-3000 m
(Ethiopia), 1 100-2700 m (East Africa).

Sardinia; Serbia, Macedonia, Albania, Bulgaria; Saudi Arabia (Jebel
Fayfa); E. Sudan, Ethiopia (Eritraea to Tigray and Beghemder;
Harar); E. Uganda, SW Kenya, N. Tanzania.

H. annulatum is very closely related to 17. H. montanum (q.v. for
differences), being its sister species and the basal species of a group
comprising the remaining species in sect. Adenosepalum (Spp. 19-
24). It varies in pubescence and glandularity over its widely disjunct
areas of distribution. The European populations have pubescent
stems and leaves and vary in the length of sepal cilia, and they lack
superficial or laminar black glands. The northern Ethiopian, Suda-
nese and Arabian populations show parallel trends north-eastward
from Tigray to Arabia in pubescence (sparsely pubescent to glabrous
stems and leaves), glandularity (black glands on stem, sepals and
petals absent or present) and sepal cilia (long to short). The southern
(Harar) plants tend towards the East African population in density of
pubescence, in the presence of occasional black glands on the
leaves, and (rarely) in having red tinges on the petal veins, but are
otherwise like the northern group. The East African population
varies in pubescence (stems and leaves densely pubescent to gla-
brous) and glandularity (black or red glands always on stems and
petals, sometimes on leaves and sepals), but the sepal cilia are
always long and the petals always red-tinged. The overlaps in
variation preclude the recognition of any of these populations at
specific level, but three variable subspecies can be distinguished.
Subsp. annulatum comprises the European populations, subsp.
intermedium most of the north-east African and Arabian populations
and subsp. afromontanum the Harar and east African populations.
For a key to these subspecies, see p. 172.

18a. Hypericum annulatum subsp. intermedium (Steud. ex A.
Rich.) N. Robson in Bull. not. Hist. Mus. Land. (Bot.) 23: 69
(1993). Type as for H. intermedium Steud. ex A. Rich.

Map 39.

H. intermedium Steud. ex A. Rich., Tent. fl. abyss. 1: 95 (1847);
Oliv., Fl. Trop. Afr. 1: 155 (1868) pro parte quoad typum; Engler
in Phys. Abh. K. Akad. Wiss. Berlin 1891: 306 (1892);
Hochgebirgsfl. Trop. Afr.: 306 (1892); T.C.E. Fr. in Notizbl. Bot.
Gart. Berlin 8: 566 (1923). Types: Ethiopia, ad latus boreale
montium altiorum prope Adoam, 20 November 1838 (fl),
Schimper II 1062 (P-lectotype; BR, Fl, G, K!, LE, MO-
isolectotypes); without precise locality or date, Quartin-Dillon &
Petit 37 (P-syntype).

H. annulatum sensu Cufod. in Bull. Jard. hot. Brux. 29, Suppl.: 588

200

N.K.B. ROBSON

Fig. 29 A. H. lanuginosum: (a) habit; (b) leaf; (c) bract; (d) sepal; (e) petal; (f) capsule. B. H. cuisinii: (g) habit. C. H. annulatum subsp. afromontanum:
(h) habit; (i) leaf; (j) bract; (k) sepal; (1) petal; (m) capsule (a, g, h x !/2; b, i x 1 ; c-f, j-m x 3). A. Bornmiiller 1 1525. B. Schlieben 5062. C. Guiol 2295.

STUDIES IN HYPER/CUM

201

Map 39 Sect. 27: 18. H. annulatum: a. subsp. intermedium specimens
•, records D; c. subsp. afromontanum specimens •, records O.

(1959); Moggi & Pisacchi in Webbia 22: 272, f. 12 (1967);
Collen., Fls Saudi Arabia: 261 & photograph (1985).
H. intermedium forma obtusifolium R. Keller ex Moggi & Pisacchi
in Webbia 22: 272 (1967), in synon.

Icon: Moggi & Pisacchi in Webbia 22: 272, f. 12 (1967).

Stem without or rarely with few black glands, sparsely puberulous to
glabrous. Leaves without laminar black glands, ± sparsely and very
shortly pubescent to puberulous or glabrous. Sepals short- to long-
glandular-ciliate (i.e. cilia shorter to 2 or more times as long as
glands), occasionally with some laminar glands black. Petals with
few (rarely more numerous) punctiform laminar black glands, rarely
red-veined in bud.

Saudi Arabia (Asir), Sudan Republic (S. Red Sea Hills), N. Ethiopia
(Eritraea to L. Tana and N. Shoa).

SAUDI ARABIA. Asir: Jabal Fayfa, 100 km NE of Jizan, 1500 m, 10
April 1982 (fl), Collenette 3536 (E, K); loc. cit., 1725 m, 28 March 1988 (fl),
Collenette 6625 (E*, K); loc. cit., 14 April 1982 (fl), Grainger 578 (E).

SUDAN REPUBLIC. Kassala, Red Sea Hills, Diris Pass, 1700 m, 10
April 1953 (fl), Jackson 2867 (K).

ETHIOPIA. Eritraea: sul monte Matara, 20 September 1902 (fl), Pappi
905 (BM, K); Asmara to Massawa, c. 1 7 km, 1 800 m, 5 February 1 969 (fl &
fr), De Wilde 4565 (K); Ad Teclesan, 2 1 00 m, 20 March 1 944 (fl), Bally 6660
(K).Tigray: vomSchollodabeiAdoa,2010m, 15 August 1862(fr),5c/»'m/w
37 (BM, E*); prope Adowa, 1 March 1837 (fl), Quartin-Dillon & Petit s.n.
(K). Shoa: Alia Amba, October 1842 (1. fl), Roth 193 (K). Begemdir: Simien
Mts and Gondar (Moggi & Pisacchi, 1967).

Subsp. intermedium varies north-eastward by decreasing pubes-
cence and increasing glandularity, so that the Sudanese and Arabian
plants are glabrous with black-gland-dotted sepals and stems. The
Arabian plants also have sepals with short glandular cilia, whereas in
the others they are medium to long. The eastern Ethiopian popula-

Map 40 Sect. 27: 18. H. annulatum: b. subsp. annulatum • (see also
Map 41); 24. H. decaisneanumA..

tion has more in common with the East African plants and has
therefore been included in subsp. afromontanum.

1 8b. Hypericum annulatum subsp. annulatum
Maps 40, 41.

H. atomarium sensu Velen., Fl. Bulg.: 105 (1891); Stjep.-Vesel. in
Josifovtf, FL Srbije 3: 1 12, t. 32 ff. 2, 2a (1972).

H. perfoliatum van annulatum (Moris) Fiori in Fiori & Paoletti, Fl.
Italia 1: 389 (1898), Nuov. Fl. Italia 1: 524 (1924).

H. degenii Bornm. in Magyar Bot. Lap. 9: 90 (1910); Stefanoff in
God. Agr.-les. Fak. Univ. Sofiya 10: 58, tt. 2 f. 20, 3 f. 34 (1932),
11: 165 (1933), 12: 85 (1934), in Pflanzenareale 4(1): Karte 4a
(1933); Jordanoff & Kozuh. in Jordanoff, Fl. R. P. Bulgaricae 4:
239, t. 45 f. 1 (excl. 6) (1970). Types: Bornmuller cited nine
syntype specimens, four from Serbia and five from Bulgaria. As
there is no apparent reason for choosing any particular specimen
as lectotype, I have selected the most widely distributed collec-
tion: Bulgaria, in rupestribus ad Stanimaka, June 1894 (1. fl),
Sth'brny s.n. (BPMectotype; BM!, FR!, K!).

H. atomarium subsp. degenii (Bornm.) Hayek, Prodr. fl. pen. Bale.
1: 536(1925).

Icones: Moris, Fl. Sardoa 1: t. 22 (1 837); Chiappini, Fl. paesag. veg.
Sardegna: t. 1 1 (1985); Jordanoff, Fl. R. P. Bulgaricae 4: t. 45 f. 1
(excl. 5) (1970).

Stem without red or black glands, densely short-pubescent. Leaves
without laminar black glands, densely short-pubescent. Sepals long-
to short-glandular-ciliate (i.e. cilia shorter than to 2 or more times as
long as glands), with laminar glands all pale. Petals without puncti-
form laminar black glands, not tinged red in bud. 2n = 1 6 (see p. 1 99).

SARDINIA. Monte Santa Vittoria di Esterzili, 1 2 1 2 m, 25 August 1 965
(fr), Bongoni 556 (BM); Nodu 'e Littipori (fide Arrigoni et al., 1973).

SERBIA. Prope Nis, June 1894 (fl), Adamovic s.n. (FR, K); prope
Alpa£kavica, 17 June 1895 (fl), Adamovic s.n. (K); ad Markovo Kale sub
monte Kostilovitza prope Vranjos, June 1896 (fl), Adamovic s.n. (K).

MACEDONIA. Treska gorge, c. 400 m, 8 June 1937 (fl), Rev. <£ Mrs
H.P. Thompson 877 (K).

ALBANIA. Shkodra: Bertiscus, in valle rivuli Locanska Bistrica, c. 700
m, 17-19 July 1933 (fr), K.H. Rechinger & Scheffer 1036 (BM, K).

BULGARIA. Sofiya: circa Dragoman, July 1 896 (fl), Jovanitz s.n. (K).
Blagoevgrad [Gorna Dzhumavo): Melnik, Tal zwischen Melnik und Roien-
Kloster, 13 June 1971 (fl), F.K. & J. Meyer 10235 (BM, JE). Plovdiv: infra
pagum Ba6kovo, oppidum Asenovgrad [Stanimaka] versus, 300 m, 12 June
1973 (fl), Greuter 1 1 175 (H); ad Papozli, June 1910 (fl & fr), Stribrny s.n.
(H); Asenovgrad, Westhange entlang das Flusses oberhalb Batschkovo-

202

N.K.B. ROBSON

Kloster, 24 June 196 1 (fl), Bisse & Schneider 376 (JE), 377 (BM). Khaskovo:
near Boju, valley to Daridere, 17 July 1926 (fl), Turrill 1444 (K).

SWITZERLAND (subspontaneous). Zurich: Stafa,430m, 12 July 1981
(fl&fr), Kramer 1500 (BM).

CULTIVATED. Specimens seen from England (1929, herbarium and
1994, living) and Sweden (1973).

Subsp. annulatum varies somewhat in leaf width, but neither this
character nor the sepal cilia length (shorter than or equalling gland in
Sardinia, usually exceeding gland by up to c. 6 times in Balkans) can
be used to separate these geographically widely separate populations.

18c. Hypericum annulatum subsp. afromontanum (Bullock) N.

Robson in Bull. nat. Hist. Mus. Land. (Bot.) 23: 69 (1993).

Type as for H. afromontanum Bullock.
Fig. 29C, Map 39.

H. afromontanum Bullock in Kew Bull. 1932: 492 (1932); Hedberg
in Symb. hot. Upsal. 15(1): 131 (1957); Agnew, Upland Kenya
wildfls: 186 (1974). Type: Kenya, Mt Elgon, 3540 m, December
1930 (fl), Major E.J. & Mrs C. Lugard 338a (K!-holotype; BM!
'338').

H. annulatum sensu Milne-Redh. in Kew Bull. 8: 435 (1953); Moggi
& Pisacchi in Webbia 22: 272, f. 13 (1967) pro parte; Agnew,
Upland Kenya wildfls: 1 86 ( 1 974); N. Robson in Bamps, Robson
& Verdcourt, Fl. trap. E. Afr., Guttif.: 30 (1978).

Icon: Agnew, Upland Kenya wildfls: 185 (1974).

Stem usually with numerous black (or very rarely red) glands,
densely to sparsely puberulous or rarely glabrous. Leaves some-
times with few to numerous laminar black glands, puberulous above
and densely pubescent beneath or very rarely wholly glabrous.
Sepals long-glandular-ciliate (i.e. cilia more than twice as long as
glands), usually with some or all laminar glands black. Petals with
few distal or numerous scattered punctiform laminar black glands,
always? red-tinged in bud.

SE Ethiopia (Harar), East Africa (eastern Uganda, south-western
Kenya, northern Tanzania).

ETHIOPIA. Harar: Gara-muleta, 2550 m, 23 October 1960 (fl),
l.E.C.A.M.A. J-54 (K); Gara Mullata mtn, ± 3060 m, 24 November 1962 (fr),
Burger 2394 (K).

UGANDA. Eastern: Mbale Distr., Mt Elgon, Bupota [Bupoto], 1500 m,
7 August 1917 (fl & fr), Snowden 522 (BM, K); Bugishu Distr., Mt Elgon, W.
slope above Butaderi, 3500 m, 5 December 1967 (fl), Hedberg 4488 (K).

KENYA. Northern Frontier: Mathews Range, Dunyus, 2 100 m, 25 June
1 944 (fl & fr). Bally 36 1 0 (K); 24 km N. of Maralal on road to Baragoi, 2 1 80
m, 26 October 1978 (fr), Gilbert et al. 5156 (K). Turkana: West Suk Distr.,
Kitale to Moroto, c. Km 64, 4 October 1952 (1. fl), Verdcourt 747 (K). Rift
Valley: Naivasha Distr., Longonot, 2700 m, March 1922 (fl), Dummer 5133
(K); Ravine Distr., 2nd day's march from Eldama Ravine, November 1898
(fl), Whyte s.n. (K). Central: Kitui Distr., Galunka, 30 May 1902 (1. fl & fr),
Kassner 873 (BM); Machakos Distr., c. 5.6 km N. of Nunguni, Kilungu
Location, 1800 m, 1 1 June 1967 (fl), Mwangangi 58 (K); Meru Distr.?, Mt
Kenia septentrionalis inter flumina Liki et Kongoni, 1 2 February 1922 (e. fr),
R..E. & T.C.E. Fries 1488 (K). Masai: Ngong Hills, S. slopes above Magadi
road, 5 August 1957 (fl), Greenway 9221 (K); Kilmandsharo, N. Seite,
Loiokitok [Laiokitok], 1850 m, 1 1 April 1934 (fr), Schlieben 5062 (BM, K,
Z).

TANZANIA. Northern: Masai Distr, Ol Doinyo Loldadwenya, 3000 m,
27 January 1962 (fl), Newbould 5903 (K); Arusha Distr., Songe Hill, near
Telegraph Hill, 201 1 m, 23 February 1969 (fl & e. fr), Richards 28184 (K);
Mbulu Distr., Shesheda to S. of Gendabi, Mt Hanang, 2250 m, 10 February
1946 (fl), Greenway 7678 (K).Tanga: W. Usambaras, Gologolo- Mkumbala
footpath, 1800 m, 4 June 1953 (fl), Drummond & Hemsley 2866 (K);
Hambalai Scarp, 1800 m, January 1933 (fl), Mr & Mrs Moreau 45 (K).

Subsp. afromontanum is characterized by an increase of black

glands (stem, leaves, sepals, petals), constant long sepalline cilia and
red-tinged petals, and the almost constant occurrence of indumentum
on stem and leaves. In these characters the Harar population from
Ethiopia agrees with the East African plants except for the absence
of black glands on the stem.

The type of H. afromontanum belongs to an upland form with a
condensed inflorescence; but this cannot be separated from the
lower-altitude form with the typical, widely spreading lax inflores-
cence on account of numerous intermediate specimens.

19. Hypericum delphicum Boiss. & Heldr. in Boiss., Diagn. pi.
orient. II, 1: 106 (1853); Boiss., Fl. orient. 1: 807 (1867);
Halacsy, Consp.fl. graec. 1: 282 (1900); Hayek, Prodr.fl. pen.
Bale. 1: 535 ( 1 925); R. Keller in Engl. & Prantl, Nat. Pflanzenfam.
2nd ed. 21: 179 (1925); Turrill in Kew Bull. 1932: 248 (1932);
Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: 58, t. 3 f. 3
(1932), 11: 164 (1933), 12: 85 (1934), in Pflanzenareale 4(1):
Karte 4a (1933); Rech. f. in Beih. Bot. Centralbl. 54B: 617
(1936), FL aegaea: 264 (1943); N. Robson in Tutin et al., Flora
Europaea 2: 265 (1968); Polunin, Fls Greece & Balkans: 337
(1980); Greuter, Burdet & Long, Med-Checklist 3: 265 (1986);
N. Robson in Cullen et al., Eur. Gdn Fl. 4: 73 (1995). Type:
Greece, Evvoia, 'prope Steni ad radices montis Delphi Eubeae',
450 m, August 1848 (e. fr), Heldreich s.n. (G!-holotype; BM!,
H!,K!).

Map 41.

Perennial herb 0. 1 1-0.35(-0.45) m tall, erect to ascending from
creeping, rooting and branching base, with herbaceous taproot,
usually several-stemmed, unbranched below inflorescence, strigose-
pubescent on stems below inflorescence and on both surfaces of
leaves. Stems green, terete; internodes exceeding to shorter than
leaves, eglandular. Leaves sessile; lamina 12-35 x 9-29 mm, ob-
long-ovate to broadly ovate, concolorous, thinly chartaceous, not
glaucous, equally and rather sparsely hirsute on both sides, more
densely so along veins beneath, plane, spreading; apex rounded,
margin entire, base rounded to shallowly cordate; venation: 3^
pairs of laterals curved-ascending from lower 0.25-0.4 of midrib,
tertiary reticulation dense, slightly prominent; laminar glands pale,
dense, subequal; intramarginal glands black, dense to rather sparse.
Inflorescence 4-c. 90-flowered from 1-3 nodes, without flowering
branches below, shortly cylindric to corymbiform, dense; pedicels
1.5-2 mm; bracts and bracteoles lanceolate to linear-lanceolate,
black-glandular-ciliate, densely glandular-auriculate. Flowers 12-
15(-20?) mm in diam.; buds ellipsoid-subglobose, rounded. Sepals
3-6 x 1-1.5 mm, equal, free, narrowly oblong to narrowly elliptic,

STUDIES IN HYPERICUM

acute to subacute, with margin long-glandular-ciliate; veins 3, promi-
nent, outer ones branching; laminar glands pale, striiform to
punctiform; marginal glands black, flat-topped. Petals rather pale
yellow, 7-9 x 2.5-3 mm, 2.5-3 x sepals, oblong-elliptic, rounded,
apiculus absent; laminar glands pale, sometimes few; marginal
glands distal, black, immersed or ± prominent. Stamens c. 20-35,
longest 7-9 mm, equalling petals; anther gland black. Ovary c. 2 x
1 mm, ellipsoid; styles 5-6 mm, 2.5-3 x ovary, spreading-incurved.
Capsule 4-5 x 2.5-3 mm, broadly ellipsoid, shorter than to exceed-
ing sepals, enclosed when developing by petals twisting together.
Seeds dark brown, 0.6 mm long; testa finely reticulate-scalariform.
2n= 16(Reynaud, 1980).

Damp and shaded places among rocks; 300-1700 m.

Greece (Evvoia, Andhros).

GREECE. Evvoia: Distr. Halkis, village Steni (Dirfis), 450 m, 4 June
1969 (fl), Stamatiadou 6465 (ATH*, BM); in monte Xerowuni [Xirovouni],
ad pagum Steni, c. 600-800 m, 13-17 July 1932 (1. fl), K.H. Rechinger26l4
(BM). Platana prope Ky mi, 25 August-8 September 1 966 (fr), K.H. Rechinger
3785 (W); valley SW of Mt Ophir, 25 August 1981 (e. fr),Akeroyd & Preston
1381 (BM, CGE*). Andhros: Panagia, Panachrantos, 30 October 1939 (fr),
Davis 1020 (BM, K); between village Falika and monastery Panahrantou,
300-500 m, 1 1 June 1969 (fl), Stamatiadou 6581 (ATH*, BM).

H. delphicum is clearly a derivative of 18a. H. annulatum subsp.
annulatum, differing from it in the shorter, creeping and branching
stems, the rougher indumentum and (apart from the high-altitude
form of subsp. afromontanum) the more condensed inflorescence. It
has a relict distribution, being restricted to isolated areas in Evvoia
and Andhros, and, in turn, is related (ancestral?) to 20. H. athoum.

20. Hypericum athoum Boiss. & Orph. in Boiss., Fl. orient. 1: 794
(1867); Hayek, Prodr.fl. pen. Bale. 1: 535 (1925); Stefanoff in
God. Agr.-les. Fak. Univ. Sofiya 10: 85, t. 3 f. 32 (1932), 11: 164
(1933), 12: 84 (1934), in Pflanzenareale 4(1): Karte 4a (1933);
Ade & Rech. f. inRepert. Nov. Sp., Beih. 100: 122 (1938); Rech.
f., Fl. aegaea: 264 (1943); N. Robson in Tutin et al., Fl.Euro-
paea 2: 265 (1968); N. Robson & Strid in Strid, Mtn Fl. Greece
1: 602,f. 35.6 ( 1986); Greuter, Burdet&Long,Med-Checklist3:
264 (1986); N. Robson in Cullen et al., Eur. Gdn Fl. 4: 73
(1995); non H. montanum var. athoum Griseb., Spic.fl. rumel.:
224 (1843). Type: Greece, Makhedonia, in fissuris rupium
regionis superioris mentis Athos Macedoniae, n.d. (fl), Orphan-
ides 240 (G!- holotype).

Map 41.

H. sanctum Degen in Oesterr. Bot. Zeitschr. 41: 333 (1891); R.
Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 178
(1925). Type: Greece, Thraki, Samothraki, in saxosis umbrosis
ad torrentes montis Phengari [Pengari], 800 m, 28 June 1890 (1.
fl), Degen s.n. (BP-holotype; K!, W!-isotypes).

203

Map 41 Sect. 27: 18. H. annulatum: b. subsp. annulatum (part) • (see
also Map 40); 19. H. delphicum D; 20. H. athoum •; 21. H. atomarium
(part) A; 22. H. cuisinii A.

Perennial herb with stems 0. 1 -0.25 m long, weak, procumbent from
creeping, rooting and branching base, with herbaceous taproot,
many-stemmed, unbranched and densely to rather sparsely pilose
below inflorescence. Stems green, terete; internodes exceeding leaves,
eglandular. Leaves with petiole 0.5-1.5 mm; lamina 8-15 x 5-10
mm, broadly ovate or suborbicular to rather broadly elliptic,
concolorous, thinly chartaceous, not glaucous, softly patent-pubes-
cent above, villous beneath, more densely so along veins, plane,
spreading; apex rounded, margin entire, base broadly cuneate to
truncate or subcordate; venation: 3 pairs of laterals curved-ascend-
ing from lower 0.35-0.4 of midrib, tertiary reticulation rather dense,
obscure, not prominent; laminar glands pale, rather dense (but
absent near midrib), subequal; intramarginal glands black, rather
dense to sparse. Inflorescence (l)2-7(-l l)-flowered from terminal
node, without flowering branches below, V-shaped, rather dense;
pedicels 1-2.5 mm; bracts and bracteoles lanceolate, black-glandu-
lar-ciliate, at least lowermost pair rather densely glandular-auriculate.
Flowers c. 10 mm in diam.; buds ellipsoid, rounded. Sepals 3.5^.5
x 0.8-1 .2 mm, equal, free or shortly united, lanceolate to narrowly
elliptic, acute, with margin long-glandular-ciliate; veins 3, outer
ones sometimes branching; laminar glands pale, striiform to puncti-
form; marginal glands black, flat-topped. Petals rather pale yellow,
faintly veined red, 7-9.5 x c. 2 mm, 2-2.5 x sepals, elliptic, obtuse,
apiculus absent; laminar glands absent; marginal glands subapical,
black, few, sessile or on short cilia. Stamens c. 25, longest 5-6 mm,
c. 0.7 x petals; anther gland black. Ovary 2.5 x 1.5 mm, ellipsoid;
styles c. 5 mm, 2 x ovary, spreading. Capsule c. 4.5 x 2.5 mm,
ellipsoid, about equalling sepals, enclosed when developing by
petals twisting together. Seeds dark red-brown, 0.6 mm long; testa
shal lowly and finely reticulate-scalariform.

Rock crevices and rocky places in shade, on limestone (marble) or
gneiss; 700-2000 m.

Greece (Athos Peninsula, Thasos, Samothraki).

GREECE. Makhedonia: Ayion Oros, Mt. Athos (see type); Kavalla, in
monte Pangaeon (Purner-Dagh), 1 600 m, 26-27 June \936(f\). K.H. Rechinger

204

N.K.B. ROBSON

Map 42 Sect. 27: 2 1 . //. atomarium (part) • (see also Map 41 ); 23. //. lanuginosum (part) • (see also Map 43).

10247 (BM, K);Thasos, Mt. Ipsarion, W. end of valley above Potamia, c. 850
m,21 July 1927 (fl),A£mmMPrertw2 584 (BM,CGE*).Thraki: Samothraki,
above Ano Kariotai, 700 m, 27 July 1 980 (1. fl & fr), Akeroyd & Preston 89 1
(BM, CGE*).

H. athoum is closely related to 19. H. delphicum, differing from it in
the spreading, more delicate habit, smaller petiolate leaves, softer
indumentum and smaller- and fewer- flowered inflorescence. It has a
similarly disjunct distribution relative to 18. H. annulatum subsp.
annulatum, but in the north Aegean region.

21. Hypericum atomarium Boiss., Diagn. pi. orient. I, 8: 114
(1849), Fl. orient. 1: 808 (1867); Halacsy, Consp. fl. grace. 1:
282 (1900); Bornm. inMitt. Thiir. Bot. Ver. 24: 26 (1908); Hayek,
Prodr. fl. pen. Bale. 1: 535 (1925) pro parte quoad subsp. eu-
atomarium Hayek; R. Keller in Engl. & Prantl, Nat. Pflanzenfam.
2nd ed. 21: 179 (1925) pro parte; Rech. f. in Ann. Naturhist.
Mus. Wien 43: 305 (1929); Stefanoff in God. Agr.-les. Fak. Univ.
Sofiya 11: 164 (1933), 12: 85 (1934) pro parte, e\c\. syn. H.
millepunctatum Holmboe praeter spec. Sintenis 602, in
Pflanzenareale 4(1): Karte 4a (1933); Rech. f., Fl. aegaea: 246
: (1943); N. Robson in Notes R.B.G. Edinb. 27: 196 (1967), in
Davis, Fl. Turkey 2: 387 (1967), in Tutin et al., Fl. Europaea 2:
266 (1968); Greuter, Burdet & Long, Med-Checklist 3: 264
(1986); Carlstrom, Surv. Fl. Phytogeogr. Rodhos, (etc.).: 63,
map 309 (1987); N. Robson in Cullen et al., Eur. Gdn Fl. 4: 72
(1995). Type: Turkey, Manisa or Izmir, in dumosis montis Sipyli
supra Magnesium [Manisa] in Lydia, June 1842 (fl), Boissier
s.n. (G!-holotype).

Maps 4 1,42.

H. hirsutum sensu Sibth. & Sm., Fl. Graecae Prodr. 2: 117(1813);

Chaub. & Bory, Exped. sci. Moree 3: 152 (1832), Nouv. fl.

Pelopp.: 53(1838).
H. lanuginosum sensu d'Urv., Enum. pis Ponti-Eux.: 58 (1822);

Friedr., Reise Neu-Griechenland: 273 (1838).

H. supinum Vis. inAtti Riun. Sci. ital.: 175 (1841), ///. piante Grec.
Asia minore: 17 (1842); Mabberley in Taxon 31: 71 (1982), pro
min. parte omnes. Types: Turkey, Balikesir?, circa Antandro ad
sinum Golfo 'Adramitti [Edremit] dictum, 1819 (fl), Parolini &

STUDIES IN HYPER1CUM

205

Webb s.n. (PAD!-syntype). The other syntype belongs to H.

tomentosum L. van (5; see Robson (1967a: 196), where it was

chosen as lectotype of H. supinum Vis.
H. lanuginosum subsp. atomarium (Boiss.) Holub (description not

traced).

Perennial herb 0.2-0.8 m tall, erect or decumbent but not rooting at
base, with ± woody taproot, few-stemmed, unbranched below inflo-
rescence or with short axillary shoots after fruit ripens, shortly
whitish-pubescent on stems below inflorescence and on both leaf
surfaces. Stems green, terete; internodes all shorter than leaves or
upper (or more rarely all) exceeding them. Leaves sessile; lamina
15^45(-55) x 8-20(-22) mm, ovate to oblong or elliptic, paler
beneath, thinly chartaceous, not glaucous, shortly pubescent be-
neath, usually shorter or puberulous above, plane, spreading; apex
rounded, margin entire, base cordate-amplexicaul to rounded; vena-
tion: 3 pairs of laterals curved-ascending from lower 0.2-0.25 of
midrib, tertiary reticulation dense, not or scarcely prominent; laminar
glands all pale or sometimes scattered black ones distally and
towards margin, dense to rather sparse, ± unequal; intramarginal
glands black, dense to rather sparse. Inflorescence ( 1 2-) 1 5-c. 200-
flowered from (2-)5-8 nodes, without flowering branches from
lower nodes, cylindric to rarely shortly and broadly rounded-py-
ramidal; pedicels 2-3 mm; bracts and bracteoles triangular-lanceolate
to linear, black-glandular-ciliate, often with basal cilia somewhat
longer but not auriculate. Flowers 15-20 mm in diam.; buds cylin-
dric, rounded. Sepals 3.5-5 x 1-2.2 mm, subequal, free or almost so,
± narrowly oblong or oblong-lanceolate to elliptic or rarely lanceo-
late to ovate, obtuse (rarely acute) to rounded, with margin
long-glandular-ciliate; veins 3, outer sometimes branched; laminar
glands pale, striiform; marginal glands black, flat-topped. Petals
pale yellow, not tinged red, (8-)9-12 x 3-3.5 mm, c. 2.5 x sepals,
elliptic to oblanceolate, rounded, apiculus absent; laminar glands
pale, punctiform to striiform, and also usually black, punctiform,
scattered; marginal glands absent. Stamens 25-40, longest 6-8 mm,
c. 0.65 x petals; anther gland black. Ovary 1.5-3 x 1-1.5 mm,
ellipsoid; styles 3.5-5.5 mm, 1.9-2.3 x ovary, spreading-incurved.
Capsule 5x3 mm, broadly ellipsoid, equalling or exceeding sepals,
enclosed when developing by petals twisting together. Seeds dark
reddish brown, 0.5-0.6 mm long; testa finely scalariform. 2n = 16
(Reynaud, 1973, 1980, 1981).

Stony places (schist or limestone) near streams or in damp shade;
30-1000 m.

Greece (E. Peloponnisos: Lakonia, Argolis; E. Aegean islands:
Lesvos, Khios, Ikaria, Samos, Rhodos), Turkey (W. Anatolia from
Bursa to Antalya). Naturalized in Portugal (cf. Ramos, 1993: 184).

GREECE. Lakonia: Mt.Taygete [Taiyetos], 1975 (fl), Contandriopoulos
75-259 (MARS). Argolis: Poros, prope Poros, 1 June 1867 (fl), Heldreich
252 (BM, K). Lesvos: Mytilini (Lesbos), ad Philia, c. 300 m, 1 8-24 May
1934 (fl), K.H. Rechinger5899(BM, K). Khios: Amadhes,31 May 1939 (fl),
Plan 233 (K). Ikaria: village Hristostomos, 300-350 m, 28 May 1970 (fl),
Stamatiadou9\Q6 (ATH*, BM). Samos: Vathy, 16-23 June 1932 (1. fl), K.H.
Rechinger 1 897 (BM). Rhodos: below Salakos, 300-350 m, 22 October 198 1
(e. fr), Davis 67985 (E); from Rhodes to Afantou, 27 May 1971 (fl),
Fagersten s.n. (H).

TURKEY. Bursa: prope Brussam, May 1874 (fl), Pichler 72 (K).
Balikesir: Ak-Tchai, ad Seitinly, 1 1 June 1 883 (fl), Sintenis 602b (E); Mt. Ida
[Kaz dag], prope Kareikos, 1 9 July 1 883 (fl), Sintenis 602 (BM, E, FR, G, JE,
K, U). Manisa: Salikli, Banya, 250 m, 10 June 1935 (fl), Wall s.n. (S); Kula,
700m, 21 June 1965 (fl), Coode & Jones 28 1 6 (E). U§ak: Ouchak (Phrygie),
910 m, 15 July 1857 (fl), Balansa 1 158 (BM, E, FR, G, JE, K, U). Izmir:
Selcuk, Efes [Ephesus], 3 May 1972 (fl), E. & G. Sezik 262 (BM); Odemis
distr., Bozdag, 1300m, 16August 1950(1. f\)Davis 18230(E). Aydin: Soke

to Mila§, 25 May 1962 (fl), Dudley D. 34994 (K); Camlik - Selcuk, 6 km, 30
m, 22 June 1954 (fl), Huber-Morath 12269 (G). Mugla: Sandras dag near
Agla, 600 m, 25 July 1 947 (fr),Davis 1 3638 (E, K); Datc.a to Marmaris, 6 km
from Emi9k, 1 10 m, 7 June 1962 (fl), Dudley D. 35448 (K). Denizli: Boz dag
near Geyran Yayla, 16 July 1947 (fl), Davis 13347 (E, K); Babadag, above
Kadikoy, 900 m, 8 June 1951 (e. fr), Davis 18426 (E), 18426A (K). Antalya:
from Alanya to Gazipa§a, 15 km, 3 June 1973 (fl), Himmetofelu 580 (BM).

H. atomarium is very closely related to 1 8a. H. annulatum subsp.
annulatum, differing from it essentially in lacking proper auricles on
the bracts and bracteoles and in having laminar black glands on the
sepals, usually on the petals and sometimes on the leaves. This last
character also distinguishes it from its sister species, 23. H.
lanuginosum, in which the sepals are usually more obtuse with
shorter glandular cilia. H. atomarium and H. lanuginosum together
form a south-eastern development from the Balkan H. annulatum, a
population which split into a western and an eastern species. The
laminar black gland character separates them well; although there
are relatively few black glands on the sepals of the 'outlying' Antalya
specimen cited above, they are numerous on the leaves. However,
variability towards the western end of the distribution of H.
lanuginosum (q.v.) may well indicate introgression from H.
atomarium.

See 22. H. cuisinii for a discussion of the relationships of that
species to H. atomarium.

22. Hypericum cuisinii Barbey in Bull. Soc. vaud. Sci. not. 21: 220
(1886),[<cM/5m/'];Boiss.,F/.onVm.,Suppl.:127(1888);Hayek,
Prodr.fl. pen. Bale. 1: 535 (1925); Stefanoff in God. Agr.-les.
Fak. Univ. Sofiya 10: 58, t. 3 f. 28 (1932), 12: 84 (1934), in
Pflanzenareale 4(1): Karte 4a (1933); Rech. f., Fl. Aegaea: 264
(1943); N. Robson in P. Davis, Fl. Turkey 2: 387 (1967), inTutin
et al., Fl. Europaea 2: 266 (1966); Greuter, Pfleger & Raus in
Willdenowia 13: 58 (1983); Greuter, Burdet & Long, Med-
Checklist 3: 265 (1986); Turland, Chilton & Press, Fl. Cretan
area: 93, map 675 ( 1 993). Type: Greece, Karpathos, in declivibus
lapidosis montis Elympo insula Karpathos, May 1883 (fl),
Pichler 139 (G!-holotype; E!, K!).

Fig. 29B, Map 41.

Icon: Stefani, Major & Barbey, Karpathos: t. 3 (1895).

Perennial herb, with stems 0.04-0. 15(-0.28) m long, decumbent or
diffuse-ascending, rooting, with rather woody taproot, many-
stemmed, ± caespitose, branched below inflorescence, pruinose to
shortly whitish-pubescent on stems below inflorescence and on both
surfaces of leaves or rarely wholly glabrous. Stems green, terete or
rarely (2)4-lined; internodes usually exceeding leaves, eglandular.
Leaves sessile or with petiole to 0.5 mm; lamina 2-15 x 2-10(-13)
mm, ovate or oblong or elliptic to orbicular or oblanceolate, paler
beneath, thinly chartaceous, not glaucous, shortly densely pubes-
cent to pruinose or glabrous beneath, puberulous to glabrous above,
plane or with margin recurved, spreading; apex rounded, margin
entire, base rounded to rarely cuneate; venation: 2 pairs of laterals
curved-ascending from lower 0.4 of midrib, tertiary reticulation
dense, obscure, not prominent; laminar glands all pale or rarely 1-3
distal black, dense to rather sparse, unequal; intramarginal or sub-
marginal glands black, irregular or rather sparse. Inflorescence

206

N.K.B. ROBSON

l-7(-21)-flowered from 1-2 nodes, sometimes with flowering
branches from 1-2 lower nodes, cylindric to subcorymbiform, ±
dense; pedicels 2-3 mm; bracts and bracteoles linear-lanceolate to
linear-elliptic, black-glandular-ciliate, often with basal cilia some-
what longer but not auriculate. Flowers 8-12 mm in diam.; buds
ellipsoid, rounded. Sepals 2.5-3.5 x 1-1.5 mm, equal, free or very
shortly united, oblong or elliptic-oblong to oblanceolate, obtuse to
rounded, with margin medium- to short-glandular-ciliate; veins 3,
outer sometimes branched; laminar glands pale, punctiform and
black, 2-12, scattered, ± punctiform; marginal glands black, flat-
topped. Petals pale? yellow, not tinged red, 5-7(-8) x 2-2.5 mm, c.
2.5 x sepals, elliptic-oblong, rounded, apiculus absent; laminar
glands pale, ± punctiform, few (or sometimes none?), and usually
black, up to 9, punctiform, mostly distal; marginal glands absent.
Stamens c. 25, longest 4-5.5 mm, 0.7-0.8 x petals; anther gland
black. Ovary 1.5 x 0.8-1 mm, ellipsoid; styles 3-3.5 mm, c. 2 x
ovary, spreading-incurved. Capsule 3-4 x 2.5-3 mm, ellipsoid to
subglobose, exceeding sepals, enclosed when developing by petals
twisting together. Seeds dark reddish brown, 0.4-0.6 mm long; testa
finely foveolate-scalariform.

In rock fissures (gneiss or limestone) or near springs; (10-)500-
1400m.

Greece (Khios, Ikaria, Karpathos, Kasos), Turkey (Izmir).

GREECE. Khios: Amadhes to high up Pelinaion, 8 July 1 939 (fl), Plan
322 (K). Ikaria: Insula Ikaria, June 1933 (fl & fr), Guiol 2295 (BM, K); in
summo jugo supra pagum Hag. Kyrikos, c. 950 m, 24-35 June 1932 (fl), K.H.
Rechinger 2223 (BM, K, S. W). Karpathos: Finike, 10 m, 27 July 1950 (fr),
Davis 18086(K); M. Lastos, decl. occidentalis montis Kalolimni.c. 1 100m,
ISJune 1 935 (fl ),£.//. & F. Rechinger 8\96(K);E\ympo, 30 May 1883(fl),
Barbey 129 (K, Z). Kasos:^eGreuter, Pfleger & Raus in Willdenowia 13: 58
(1983).

TURKEY. Izmir: Boz Dag, c. 1400 m, 5 July 1968 (fl)), Sorger 68-16-
5 (BM).

H. cuisinii is clearly a derivative of 21. H. atomarium growing in
higher and more exposed habitats. Whilst H. atomarium normally
has erect, non-rooting stems, in Sintenis 602 (Kaz Dag) and
Stamatiadou 9106 (Ikaria, 300-350 m) the stems are decumbent
and, in the latter, rooting. There is, nevertheless, a distinct though
small 'gap' between the respective flower sizes of the two taxa, and
so I prefer to retain H. cuisinii at specific rank, at least until
comparative field studies have been made.

The Karpathos population is the most reduced, with leaves small,
often orbicular to oblanceolate and (along with the stems) some-
times glabrous. It shows a greater resemblance in leaf shape to the
population of//, atomariumfrom adjacent Rhodos than to the rest of
H. cuisinii and could conceivably have evolved independently - as
indeed could the population on the Anatolian mainland (Boz Dag),
found at 1400 m near a spring.

23. Hypericum lanuginosum Lam., Encycl. 4: 171 (1797); Willd.,
Sp. pi. 3: 1466 (1802); Choisy, Prodr. monogr. Hyperic.: 52
( 1 82 1 ), in DC., Prodr. 1: 55 1 ( 1 824) (excl. loc. grace.); Spach in
Annls Sci. nat. (Bot.) II, 5: 357 (1836); Boiss., Fl. orient. 1: 807
(\861);Ho\mboe\nBergensMus.Skr.U,l(2): 127J.41 (1914);
R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 179
(1925); Post, Fl. Syria 2nd ed. (ed. Dinsmore) 1: 233 (1932);
Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: 58, t. 3 f. 35
(1932), 11: 165 (1933), 12: 85 (1934), in Pflanzenareale 4(1):
Karte 4a (1933); Rech. f. in Arkivf. Bot. 5: 292 (1960); Zohary,
Fl. Palaestina 1: 223, t. 329 (1966); N. Robson in Notes R.B.G.

Edinb. 27: 196 (1967), in P. Davis, FL Turkey 2: 386 (1967), in
Meikle, Fl. Cyprus 1: 297 (1977); Mouterde, Nouv. Fl. Liban
Syrie 2: 527 (1970), t. 228 f. 2 (1986); Greuter, Burdet & Long,
Med-Checklist 3: 268 (1986); N. Robson in Cullen et al., Eur.
Gdn Fl. 4: 72 ( 1 995).Type: Lebanon?, 'Levant', ? in Herb. D. de
Jussieu (P-holotype).
Maps 42, 43.

H. pestalozzae Boiss., Diagn. pi. orient. 1,8: 113(1 849), Fl. orient.
1: 808 ( 1 867); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd
ed. 21: 179 (1925); Stefanoff in God. Agr. -les. Fak. Univ. Sofiya
11: 164 (1933), 12: 85 (1934), in Pflanzenareale 4(1): Karte 4a
(1933). Type: Turkey, Antalya, in Pamphylia prope Adalia

Map 43 Sect. 27: 23. H. lanuginosum (part) • (see also Map 42).

STUDIES IN HYPERICUM

207

[Antalya], 1846 (fl), Pestalozza s.n. (G!-holotype, El-photo-
graph).

H. scabrellum Boiss., Diagn. pi. orient. II, 5: 69 (1856), Fl. orient.
1: 808 (1867); R. Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd
ed. 21: 179 (1925); Stefanoff in God. Agr.-les. Fak. Univ. Sofiya
11: 26 (1933), 12: 85 (1934), in Pflanzenareale 4(1): Karte 4a
( 1 933). Type: Turkey, Icel, ad fontes [origines] Cydni, 1 500 m, 22
September 1853 (e. fr), Kotschy 330a (also 358m, 358n) (G!-
holotype, E!- photograph; W!).

H. gracile Boiss., Diagn. pi. orient. II, 5: 170 (1856). Type: Turkey,
I9el, ad Giilek-Boghas, au nord de Tarsous, 19 August 1858,
Balansa 370 (G!-syntype); Bouloukli, pres de Mersina, 4 June
1855 (fl), Balansa 669 (GMectotype, selected here; BM!, E!, G-
P!, K!-isolectotypes).

H. lanuginosum [var.] (3 gracile (Boiss.) Boiss., Fl. orient. 1: 808
( 1 867), Suppl.: 1 29 ( 1 888); Post, FL Syria 2nd ed. (ed. Dinsmore)
1:233(1932).

H. lanuginosum subsp. gracile (Boiss.) [Holmboe in Bergens Mus.
Skr. II, 1(2): 129 (1914) in adnot., nom. provis. ex] J. Thiebaut, Fl.
Lib.-Syr.: 140 (1936); Rech. f. inArkivf. Bot. 5: 292 (1960).

H. lanuginosum subsp. millepunctatum Holmboe in Bergens Mus.
Skr. II, 1(2): 129 (1914), excl. syn. H. supinum Vis. et spec.
Sintenis 602: Lindb. f., her Cypr.: 23 (1946). Type: Cyprus,
Kyrenia distr., Lapithos, 3-4 June 1905 (fl), Holmboe 836 (BG?-
syntype); in monte supra Lapithos, 31 May 1880 (fl), Sintenis &
Rigo 616 (W-lectotype; K!).

H. lanuginosum var. scabrellum (Boiss.) N. Robson inNotes R.B.G..
Edinb. 27: 196 (1967), in P. Davis, Fl. Turkey 2: 386 (1967).

H. lanuginosum var. pestalozzae (Boiss.) N. Robson inNotes R.B.G.
Edinb. 27: 196 (1967), in P. Davis, Fl. Turkey 2: 387 (1967).

H. atomarium sensu Osorio-Tafall & Seraphim, List vase, pis Cy-
prus: 71 (1973).

Icones: Zohary, Fl. Palestina 1: t. 329 (1966); Mouterde, Nouv. Fl.
Liban Syrie 2: t. 228 f. 2 (1986).

Perennial herb 0.12-0.8 m tall, erect to ascending or occasionally
decumbent but not rooting at base, with ± woody rootstock, few-
stemmed, unbranched below inflorescence, with stems shortly
whitish-crisped-pubescent to puberulous below inflorescence or
rarely glabrous, leaves densely whitish-pubescent to rarely
scabrellous or subglabrous on both surfaces. Stems green, terete or
uppermost internodes 2-4-lined; internodes exceeding to shorter

than leaves, eglandular. Leaves sessile; lamina 15-60 x 5-25 mm,
ovate to oblong or lanceolate, paler beneath, thinly chartaceous, not
glaucous, densely shortly pubescent or velutinous to scabrellous
beneath, more shortly so above, plane, spreading; apex obtuse or
rarely subacute to rounded, margin entire, base cordate-amplexicaul
to rounded or rarely broadly cuneate; venation: 2-4 pairs of laterals
curved-ascending from lower 0.25-0.35 of midrib, tertiary reticula-
tion rather dense, obscure, not prominent; laminar glands all pale or
usually a few black scattered distally and towards margin, dense,
unequal; intramarginal glands black, dense or rather irregular. Inflo-
rescence 5- over 150-flowered from 1-8 nodes, sometimes with
flowering branches from up to 3 lower nodes, cylindric to rounded-
pyramidal; pedicels 2-4 mm: bracts and bracteoles elliptic to
linear-lanceolate, black-glandular-ciliate and sometimes -auricu-
late. Flowers 15-20 mm in diam.; buds cylindric to ellipsoid,
rounded. Sepals (3-)4-7 x (0.8-)l-3 mm, unequal to equal, free or
shortly united, broadly to rather narrowly oblong or elliptic to
subspathulate, obtuse (or rarely subacute) to rounded, with margin
long-glandular-ciliate to glandular-denticulate or very rarely entire
and eglandular; veins 5, outer ones commissural; laminar glands
pale only, shortly striiform to punctiform; marginal glands black,
flat-topped. Petals pale yellow, rarely red-veined, 7-1 2 x 3-5 mm, c.
2 x petals, elliptic to oblong, rounded, apiculus absent; laminar
glands pale, ± elongate-punctiform, scattered, or absent, rarely also
a few subapical laminar and/or marginal black gland dots. Stamens
c. 30-55, longest c. 4-8 mm, c. 0.65 x petals; anther gland black.
Ovary 1.5-3 x 1-1.5 mm, ellipsoid; styles 2.5-5 mm, 1.5-1.65 x
ovary, spreading. Capsule 4-6 x 2-4 mm, broadly ellipsoid, en-
closed when developing by petals twisting together. Seeds blackish
red, 0.6 mm long; testa finely scalariform. 2n = 16 ( n = 8, Reynaud,
1981), 32(Reynaud, 1980).

Calcareous rocks and macchi, usually in shade or near moisture; 0-
2400 m.

Turkey (southern Anatolia from Antalya to Hatay), western Syria,
Lebanon, Israel (south to Bethlehem), Jordan (Gilead), Egypt (Si-
nai)?, Cyprus.

TURKEY. Antalya: 2 km westlich Antalya, 10-40 m, 27 May 1962 (fl),
Ehrendorfer 62-1/45-28 (WU); Takhtali dag (Kemer), near Kazdere, 1000
m, 15 August 1947 (fl), Davis 14210 (E, K). Konya: 60 km westlich Mut
(Strasse Ermenek), c. 600 m, 7 June 1966 (fl), Sorger 66-30-4 (Herb.
Sorger). I?el: Anamur, about the Anamorian, 8 May 1974 (fl), JownsendlM
62 (K); Gozne, 19 June 1971 (1. fl), Ayanoglu H21 (BM). Seyhan: Karatepe,
200m, 23 June 1971 (fl), Sorger 71-31-26 (BM, Herb. Sorger); Feke distr.,
Goksu gorge below Himmetli, 700-800 m, 9 July 1952 (e. fr), Davis, Dodds
& fetik D. 19869 (BM, E, K). Hatay: Antakya, near St Peter's Church, 150-
300 m, 27 April 1957 (fl), Davis & Hedge D.27252 (BM, K); Ilica to Arsuz,
Amanos daglari, c. 500 m, 9 June 1967 (fl), Akman 72 (ANK, BM).

SYRIA. North: in vicinitate oppidum Antakieh, in rup. calc. montis
Silpius, 24 May 1933 (e. fr), Samuelsson 5328 (K): Monts Nussairy, Ai'n
Halakim, 750-900 m, June 1910 (fl), Haradjian 3454 (K). South: Leontes
[Nahrel Litani] valley, 1863-1 864 (fr),Lowne s.n. (K); Duma, 30 June 1865
(fr), G.E. Post s.n. (K).

LEBANON. Coast: Nahr el Kelb, 15 May 1943 (fl), Davis 6079 (BM,
K); Beirut, May 1886 (fl), Post s.n. (UPS); pres de Saide, 24 May 1853 (fr),
Blanche 59 (JE, K). Lower mountains: near Ain Zehalta, 1000 m, 26 May
1977 (fl), Ball 2191 (K); Kalaleh, infra pag. Aley, c. 600 m, 1 June 1932 (fl),
Samuelsson 1952(K);Bhamdun, 1200-1300 m, lOJune l9lO(f[),Bornmiiller
1 1 525 (BM). Medium mountains: Bscherre [Bcharre], ad fontes MarsTserkis,
1350 m, 19 July 1855 (fr), Kotschy 263 (BM, K, UPS).

ISRAEL. Galilee: Mt Carmel, Wadi Shomrya, 9 April 1942 (fl), Davis
4398 (BM. K); west of Cana, 16 May 191 1 (fl), Meyer 4782 (K). Samaria:
Nablus, Mt Garazim [Gerizim], 23 April 1942 (fl), Davis 4455 (K). Judaea:
Bethlehem, 1820 (fr), Gerhard s.n. (JE); in Hierosolyma [Jerusalem], pre-
1861 (fl). Roth s.n. (K).

208

JORDAN. Gilead: Rami to Bugeia, 7 June 1942 (fr), Davis 4809 (BM,
K); Wadi Qairua, 1942? (fl), Davis 4830 (BM) - not in Map 43.

EGYPT. Sinai: Sinai, 1837 (fl), Aucher 868 (G), 869 (K). This species
has not been recorded from Sinai since 1837 and can be presumed to be
extinct there now.

CYPRUS. Kyrenia distr., Lapithos, 13 June 1969 (e. fr), Lindberg s.n.
(H, K); Kykko mts, Roudkias, 24 June 1968 (fl), Economides ARI 1239 (K).

H. lanuginosum is rather variable, and two extreme reduced forms
with stems glabrous or almost so were recognised as 'rather indefi-
nite varieties' in Flora of Turkey (Robson, \961b). Var. scabrellum,
with stems weak and decumbent, leaves scabrellous to glabrous and
sepals glandular-ciliate, occurs mostly at higher altitudes through-
out the Turkish range of the species; but there are many intermediate
forms, and this is probably not a homogeneous taxon but merely the
result of high altitude and/or exposure. Specimens of this 'variety'
include:

Antalya: Calbali dag near Fesliken yayla, 1800 m, 14 July 1949
(fl), Davis 15349 (E, K). Burdur: Burdur to Antalya, 8 km from
Bucak, 720 m, 1 1 June 1962 (fl), Dudley D.35689 (K). Icel: Bulgar
[Bolkar] dag, source of Cydnus [Pamuk], 1500 m, 22 September
1853 (e. fr), Kotschy 330a (G, W). Seyhan: mts N. of Haruniye,
ravine of Cattak Suyu, 330 m, 13 June 1953 (e. fr), Huber-Morath
1 2089 (G).

The other extreme glabrous-stemmed form, var. pestalozzae, with
leaves pubescent to puberulous and sepal margin entire and
eglandular, is endemic to a small area south-west of Antalya; but
here too there are intermediates, notably just east of Antalya at
Konya Alti. Specimens of this 'variety' include:

Antalya: Kemer distr., 0-100 m, 7 July 1 949 (fr), Davis 1 5020 (E,
K); Kara dag, 200 m, 23 June 1958 (e. fr), Little 111 (E, K).
Intermediates include: Antalya: Atbiikii Bay, 5 km N. of Cirali, 26
May 1 950 (ft), Huber-Morath 9540 (BASBG); Konya Alti, 10m, 17
May 1936 (fl), Tengwall 610 (K, S).

In view of the intermediates between both these extreme forms
and typical H. lanuginosum, it is logically impossible to recognize
them taxonomically.

24. Hypericum decaisneanum Coss. & Daveau in Bull. Sac. hot.
Fr. 36: 104 (1889); E. Durand & Barratte, Fl. libic. Prodr.: 48
(1910); Keith, Prelim. Checklist Lib. FL: 567 (1973); Ali in Ali
& Jafri, Fl. Libya, Guttiferae: 8 ( 1 976); Greuter, Burdet & Long,
Med-Checklist 3: 265 (1986). Types: Libya, Cyrenaica, in
montibus apricis Cyrenaicis ad Dernah, 15 July 1875, Daveau
s.n. (P-syntype); ad Oued Dernah, 5 May 1 877 (fl), Taubert s.n.
(P-lectotype, selected here; BM!, G, K!, Z!-isolectotypes). The
BM and K specimens are labelled 'Taubert in Barbey 582', the
others have no number.

Map 40

H. taubertii Asch. & Barbey ex Coss. in Bull. Soc. hot. Fr. 36: 104
(1889) pro syn. ['taubert?]; R. Keller in Engl. & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 179 (1925); Stefanoff in God. Agr.-les.
Fak. Univ. Sofiya 11: 163 (1933), 12: 84(1934), inPflanzenareale
4(1): Karte 4a (1933). 'Type' as for H. decaisneanum Coss. &
Daveau.

Perennial herb with stems 0.04-0.15 m tall, erect to suberect,
sometimes from decumbent and rooting base, with woody taproot,
many-stemmed, unbranched below inflorescence, whitish- to grey-
ish-pubescent except inflorescence. Stems green above, ± reddish
below, all terete or inflorescence branches 2-lined; internodes all
shorter than the crowded leaves except in inflorescence and some-
times towards base, eglandular. Leaves sessile; lamina 6-12 x
3.5-10 mm, ovate, concolorous, rather thickly chartaceous, not

N.K.B. ROBSON

glaucous, shortly pubescent above, more densely and sometimes ±
crisped-pubescent beneath, with margin recurved to revolute, apex ±
depressed-concave and veins impressed, spreading to deflexed;
apex subacute to rounded, margin entire, base broadly cuneate to
cordate-amplexicaul; venation: 3-4 pairs of laterals curved-ascend-
ing from lower 0.25-0.5 of midrib, tertiary reticulation obscure;
laminar glands pale, dense, subequal and occasionally a few black;
intramarginal glands black, rather dense, regular. Inflorescence 3-c.
20-flowered, from 1-4 nodes, without lower flowering branches,
narrowly pyramidal to subcorymbiform, all or individual parts
dense; pedicels 1-1.5 mm; bracts and bracteoles oblong to triangu-
lar-lanceolate, black-glandular-ciliate, densely glandular-auriculate.
Flowers c. 12-15 mm in diam.; buds ovoid, subacute. Sepals 2.6—4.5
x 1 mm, subequal, shortly united, narrowly oblong to broadly
elliptic, rounded, with margin rather long- to short-glandular-ciliate
or -subdenticulate; veins 5, unbranched?; laminar glands pale, linear
to punctiform, occasionally with up to 6 black dots distally; marginal
glands black, flat-topped. Petals bright yellow, veined red, 6-8 x
2.5-3 mm,c. 2.5 x sepals, oblong, rounded, apiculus absent; laminar
glands pale and/or black, scattered, punctiform; marginal glands
black, few sessile. Stamens c. 40, longest 5-7 mm, 0.8-0.9 x petals;
anther gland black. Ovary c. 1 .5-2.5 x 0.8-1 .3 mm, narrowly ovoid-
pyramidal; styles c. 3.5-6 mm, c. 2.4 x ovary, spreading-incurved.
Capsule 4-5 x c. 2.5-3 mm, ovoid, equalling or exceeding sepals,
enclosed when developing by petals twisting together. Seeds not
seen.

Crevices of limestone rock; 20-700 m.

Libya (Cyrenaica - east end of Jebel el Akhdar).

LIBYA. Cyrenaica: Wadi Derna, 8 April 1939 (o. fr), Simpson 39457a
(BM) = Sandwith 248 1 (K); Derna, 26 June 1 9 1 2 (fl & fr), Vaccari 6 1 8 (BM,
Fl*); El Gubba, 13 May 1934 (bud), Pampanini & Pichi-Sermolli N.4971
(FI*,K); Wadi Shira, above falls, 600m, 18 May 1958 (fl),Park 525 (K); near
El Minier, 450 m, 1 1 April 1939 (st), Simpson 39457b (BM).

H. decaisneanum, with its densely hairy leaves and black-gland-
dotted petals, resembles a dwarf version of 1 8c. H. annulatum subsp.
afromontanum, to which it is probably most closely related. It
bridges the gap in distribution between the European and African
subspecies of//, annulatum but is much smaller than either. Accord-
ing to Gimingham & Walton (1954), it is confined to the lower slopes
(152-238 m) of the main north-facing escarpment slope of Jebel el
Akhdar as part of a well-developed chasmophytic community.

Sect. 28. ELODES (Adans.) W. Koch, Syn. FL Germ.
Helv. 2nded. 1: 148(1843).

Perennial herb with stems up to 0.4 m or longer, with simple hairs,
with dark (reddish) glands on sepals only; branching below inflores-
cence lateral. Stems terete, eglandular; cortex not exfoliating; bark
absent. Leaves opposite, decussate, sessile, free, persistent; lamina
entire, with venation subpalmate to palmate, closed except for
lowermost veins, with tertiary venation densely reticulate; laminar

STUDIES IN HYPERICUM

209

glands punctiform; marginal gland dots dense; ventral glands ab-
sent. Inflorescence 1-13-flowered, with branching dichasial (first
node) then monochasial, from one node, without lower flowering
branches; bracts and bracteoles reduced, transitional in form to
sepals. Flowers pseudo-tubular, homostylous. Sepals 5, c. 0.3-0.5
united, persistent, erect in flower and fruit, with margin glandular-
ciliate; veins 3(5); laminar glands pale, linear; marginal glands
reddish, flat-topped. Petals 5, persistent and twisting together round
developing capsule, without apiculus; marginal and laminar glands
absent. Stamen fascicles 5, united 2+2+1 (i.e. '3'), persistent, total-
ling 12-13 stamens; filaments 0.6-0.7 united; anthers yellow, gland
amber; pollen type IX; lodicules 3, scale-like, alternating with
stamen fascicles. Ovary with 3 parietal placentae, each °°-ovulate;
styles 3, free, with bases discrete; stigmas narrowly capitate. Cap-
sule 3-valved, chartaceous, with valves narrowly longitudinally
vittate. Seeds narrowly cylindric to ellipsoid-cylindric, not carinate,
without apical expansion; testa ribbed-scalariform.

BASIC CHROMOSOME NUMBERS (x). 10 and/or 8; ploidy 2 and
possibly 4. My count (n = 16; Robson, 1968), for which the slide
preparation used has deteriorated and cannot now be checked, and
that of Al-Bermani et al. (1993) - 2n = 16, i.e. n = 8, fit better with
the morphology of H. elodes, which is specialized in relation to H.
coadunatum (n = 9) in all respects. On the other hand, Delay's count
(n = 10; Delay, 1972) is supported by a photograph and would imply
that H. elodes is sister species to the whole H. caprifolium group
(Spp. 13-15; n = 9-8). Only further counts will reveal which is
correct.

HABITAT. Wet soil, mesotrophic mires, along streams and in ponds
on acid soil, sometimes in deeper water; sea level to c. 700 m (Spain,
Azores).

DISTRIBUTION. British Isles; continental Europe from France, Spain
and Portugal eastward to NW Germany (and scattered localities
further east), Austria ? and NW Italy; Azores.

One species.

1. Hypericum elodes L., Amoen. Acad. 4: 105 (1759), Sp. pi, 2nd
ed.: 1 106 (1762); Hudson, Fl angl.: 292 (1762); Choisy, Prodr.
monogr.Hyperic.: 52 (l82l),inDC., Prodr. 1:551 (1824);Syme,
Engl. Bot. 3rd ed.: 159, t. 276 (1 863); H.Watson in Godman, Nat.
Hist. Azores: 143 (1870); Trelease in Rep. Mo. hot. Gdn 8: 100
(1897); Ohlendorf, Beitr. Anat. Biol. Fruchte u. Samen einheim.
Wasser- u. Sumpfpflanzen: 59 (1907) [Diss. Erlangen U.]; R.
Keller in Engl. & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 175
(1925); Hegi, ///. Fl. Mittel-Europa 5(1): 509, ff. 1997-8 (1925);
Stefanoff in God. Agr.-les. Fak. Univ. Sofiya 10: 58, t. 1 f. 1, t. 2
f. 1, t. 3 f. 1 (1932), 11: 139 (1933), 12: 81 (1934), in
Pflanzenareale 4(1): Karte la (1933); Mansfeld in Repert. Sp.
nov. 47: 278 (1939); Guinea, Viscaya Paisaje veg.: 215 & maps
(1949); Ross-Craig, Drawings Br. Pis 6: t. 16 (1952); Font Quer
in Geogr. Univ. de Vidal de la Blache 10: 207, f. 75 (1953); Corti
in Webbia 11: 847 (1956); Janchen in Hofler & Knoll, Cat. Fl.
Austr. 1(2): 257 (1957) ['helodes']; Pinto da Silva & Rainha in
Agronomia lusit. 20: 236 (1959); Palinha, Cat. PL Vase. Azores:
75 (1966); Bournerias mBull.Ass. Nat. Vail. Loingte: 86 (1967);
N. Robson in Tutin et al., Fl. Europaea 2: 266 (1968); Fournier,
Quatre flares France 2nd ed.: 456 (1977, repr. 1990) ['helodes'];
van Rompaey & Delvosalle, Atlas Fl. Belg. Lux. Comment.: 66
( 1 978), Atlas, 2nd ed. : t. 4 1 8 ( 1 979); Bonafe Barcelo, Fl. Mallorca
3: 178 (1979); Pignatti, Fl. Italia 1: 347 (1982); Ramos in Trab.
Dep. Bot. Univ. Complut. 12: 54, t. 6 f. 5 (1983); I. Hagemann in
Flora 173: 130, ff. 35, 38 (1983); Reynaud inAdansonia, no. 1:

88, t. 2 ff. 3, 4 (1985); N. Robson in Wild Fl. Mag. no. 403: 17
(1985); Mennema, Quene-Brot. & Plate, Atlas Nederl. Fl. 2: 176
(1985); Greuter, Burdet & Long, Med-Checklist 3: 265 (1986);
Clapham, Tutin & Moore, FL Br. Isles 3rd ed.: 116 (1987);
Ramos in Valdes, Talavera & Galiano, Fl. Vase. Andal. Occ. 1:
318 & f. & map (1987); Haeupler & Schonf., Atlas Farn- u.
Blutenpfl. Bundesrep. DeutschL: 330, Karte 963 (1988); Kaplan,
Grenzheuser & Lenski in Tuexenia N.S. No. 9: 49 ( 1 989); Perring
& Walters, Atlas Br. FL repr. amend.: 60 (1990); Stace, New FL
Br. Isles: 257 (1991); Ramos in Castroviejo et al., FL iberica 3:
1 85 ( 1 993); N. Robson in Cullen et al., Eur. Gdn FL 4: 73 ( 1 995).
Type: Ray, Syn. Meth. Stirp. Brit. 3rd ed., ed. Dillenius: 344
(1724): 'Western Parts of England; especially towards the Land's
End in Cornwal'. Herb. Sloane 124: 9 (Herb. Buddie.) (BM-SL!)
is selected here as the neotype. It has no direct connection with
Dillenius, but Dillenius studied the Buddie collection when
working on the 3rd edition of Ray's Synopsis. The specimen is
annotated with references to Ray's Historia plantarum (1686-
1688) and Petiver, Herb. Brit.: t. 60 f. 10 (1764).
Fig. 27D, Map 36.

H. palustre Salisb., Prodr. stirp. horto Chapel Allerton: 369 (1796),

nom. illegit. Type as for H. elodes L.
Elodea palustris J. St. Hil., Exp. Fam. nat. 2: 24 (1805). Type as for

H. elodes L.
Elodes palustris Spach \nAnn. Sci. nat. (Bot.) II, 5: 1 72 ( 1 836), Hist.

nat. veg. Phan. 5: 369 (1836); Rchb., Ic.fl. germ. helv. 6: t. 342

(1844); Willk. & Lange, Prodr. fl. hispan. 3: 596 (1878); Gluck,

Biol. morph. Untersuch. Wasser- u. Sumpfgewdchse 3: 38 (191 1),

nom. illegit. Type as for H. elodes L.
Hypericum tomentosum sensu Durand in Steud., Nomencl. hot. 2nd

ed. 1: 790 (1840), nom. synon.
H. helodes St.-Lager in Ann. Soc. hot. Lyon 7: 128 (1880) et auct.

plur., orth. mut.
H. helodeum St.-Lager in Ann. Soc. hot. Lyon 7: 128 (1880), orth.

mut.
Helodes glandulosum St.-Lager, Nomencl. hot.: 37 ( 1 88 1 ); Morot in

J. de Bot. 8: 84 (1894); Bubani, FL Pyren. 3: 352 (1901)

['glandulosa'], nom. illegit. Type as for Hypericum elodes L.
Tripentas helodes (L.) Asch. & Graebn., FL Nordostdeutschen

FlachL: 493 (1899), comb, illegit. (Art 63.1).
Elodes palustris forma submersa Gluck, Biol. morph. Untersuch.

Wasser- u. Sumpfgewdchse 3: 39 (1911). Type: No specimen

made?
Hypericum elodes forma glabratum Druce in Rep. Bot. Exch. Cl. 7:

435 (1924). Types: England, Hampshire (south), Lyndhurst, 1924,

Druce s.n. (OXF-syntype); Buckinghamshire, Burnham Beeches,

1922, Druce s.n. (OXF-lectotype, selected here).
Spachelodes elodes (L.)Y. Kimura in J. jap. Bot. 11: 832 (1935), in

Nakai & Honda, Nova fl. jap. 10: 19 (1951).
Hypericum helodes forma terrestre Gluck in Panscher, Susswasser-

Flora Mitteleuropas 15: 297 (1936). Type: No specimen made?

Icones: Syme, Eng. Bot. 3rd ed.: t. 276 (1863); Ross-Craig, Draw-
ings Br. Pis 6: t. 16(1952);

Perennial herb with herbaceous rootstock emitting erect (wholly
submerged) to prostrate and rooting (terrestrial) stems up to 0.4(-
0.7) m, the terrestrial or shallow-water shoots bearing erect to
ascending, terminal and axillary flowering shoots 0.05-0.2 m tall,
the whole emergent plant up to sepals (dorsal) crisped-pubescent to
tomentose or inflorescence puberulous, submerged parts shortly and
finely villous to glabrous. Stems green to reddish, terete, 1 mm thick
and threadlike (submerged) to 6 mm thick with swollen spongy

210

N.K.B. ROBSON

internodes (shallow water or terrestrial); internodes up to 50(-80)
mm long and 5 x as long as leaves (deep water) to shorter than leaves
(terrestrial). Leaves sessile; lamina 5-30 x 2-22 mm, oblong-elliptic
(deep water) to broadly elliptic or broadly ovate or orbicular,
concolorous, not glaucous, plane, spreading; apex rounded, base
cuneate (deep water) to cordate-amplexicaul; venation: 2-3 pairs of
laterals curved-ascending from base or near base of midrib; tertiary
reticulation dense, plane; laminar glands pale, dense, scattered,
small; intramarginal glands pale, dense. Inflorescence 1-13-flow-
ered from one node, terminal but usually apparently axillary when
only one of uppermost pair of axillary shoots develops, laxly
subcorymbiform to narrowly pyramidal, without lower flowering
branches; pedicels 3-7 mm (-1 1 mm in fruit); bracts not auriculate;
bracteoles triangular, obtuse, red-glandular-ciliate. Flowers 7-15
mm in diam., infundibular to subrotate, pseudo-tubular; buds cylin-
dric to narrowly ovoid, subacute. Sepals 2.5-3.5 x 1-2 mm, subequal,
c. 0.3-0.5 united, ovate or triangular-ovate to narrowly oblong,
rounded, with margin glandular-ciliate; veins 3(5), branched or not;
laminar glands pale, linear; marginal glands red, on short cilia.
Petals lemon yellow, not tinged red, 8-10 x 2.7-3 mm, c. 3 x sepals,
oblong-oblanceolate, subtruncate, without apiculus, in lower 0.25
with adnate ligulate appendage, apex free, trifid; laminar and mar-
ginal glands absent. Stamens 12-13, clearly 3-fascicled, with
filaments c. 0.65 connate with line of hairs below free parts, longest
c. 5-6.5 mm, c. 0.6 x petals; anther gland amber; lodicules 3,
squamiform, 0.5 x 0.25 mm, elliptic, retuse. Ovary 1-locular, c. 2.5-
2.7 x 1-1.5 mm, narrowly ovoid; styles 3, 2-2.5 mm, just shorter
than ovary, ± narrowly outcurving. Capsule 4-5 x 2.5-3 mm, ovoid
to cylindric, exceeding sepals. Seeds dark yellow-brown, 0.6-0.8
mm long; testa ribbed-scalariform (cf. Reynaud, 1985: 91, t. 2 ff. 3,
4). 2n = 16 (n = 8) (Al-Bermani et al., 1993), 20 (n = 10) (Delay,
1972), 32 (n = 16) (Robson, 1968); see sectional description.

In mesotrophic mires and pond or stream margins, usually in shal-
low water but sometimes in wet soil or deep water (to c. 50 cm); to
800 m in Spain and 770 m in the Azores, lowland elsewhere in
Europe.

Portugal (W. & N.), Gibraltar, Spain (N. & central), Balearic Is.
(extinct?), France (except SE), Belgium, Holland, England (except
NE), Wales, Scotland (SW and western islands), Ireland (except
centre); Azores. Unlocalized specimens in Herb. Schousboe are
likely to come from Iberia rather than Morocco (Jahand. & Maire,
Cat. Pis Maroc 3:484, 1934).

PORTUGAL. Porto: near Oporto, 2 July 1887 (fl), Murray s.n. (BM).
Viseu: Santa CombaDao, Pego, 15 June 1954(fl), J. & A. Matos & Marques
4996 (BM, COI*). Coimbra: Montemar-o-Velho, Matas de Foja, 8 August
1950 (fl & fr), J. & A. Matos s.n. (BM, COI*). Leiria: Nazare, margens da
Lagoa de Gataias, 25 July 1952 (fl & fr), A. & R. Fernandes & Sousa 4223

(BM, COI*). Santarem: prope Salvaterra dos Magos, July 1890 (fl & fr),
Daveau in Monteiro Fl. lus. exsicc. 4252 (BM). Setubal, Lagoa de Albufeira,
20 July 1941 (fr), Pedro 40 (K, LISJC*).

SPAIN. La Corufia: Puente Carreira toAyazo, 9 August 1968 (fl), Bellot
s.n. (W). Pontevedra: peninsula El Grove, 7 July 1930 (fl), Buck s.n. (H).
Lugo: Begonte, 5 August 1964 (fl), Bellot & Casaseca s.n. (M*, RNG).
Viscaya: Urquiola, 27 August 1946 (fr), Guinea 634/575 (RNG, SEV*).
Alava: entre Ochandiano et Villareal, 700 m, July 1935 (fl), Losa in Sennen
9894 (BM). ? : Aragon, au pied de la montagne de Castanede, 800 m, July
1 864 (fl), du Parquet s.n. (BM). Salamanca: between Salamanca and Ciudad
Rodrigo, 30 June 1928 (fl), Lacaita 532/28 (BM). Toledo: Monies de Toledo,
Monies de Mora, 18 June 1959 (fl), Sandwith 5579 (K). Huelva: Almonle,
Marismas de El Rocio, Laguna del Acebron, 23 June 1981 (fl), Demisse,
Silvestre & Valdes 54 (RNG).

BALEARIC ISLANDS. Mallorca, Arta, nol seen since 1908 (/WeBonafe
Barcelo, 1979: 179).

FRANCE. Finislerre: Ushanl, 1 1 Augusl 1933 (fl & fr), Meinertzhagen
s.n. (BM). Manche: Cherbourg, 30 July 1887 (fl), Corbiere s.n. (BM).
Calvados: Vire, October 1 837 (fr), Lenormand20 (K). Seine-el-Marne: Forel
de Fonlainebleau, Belle Croix, 19 Augusl 1877 (fl & fr), Moreau s.n. (H).
Yvelines: Forel de Rambouillet, aux environs de Poigny-la-Foret, 14 July
1971 (fl), Retz 64560 (BM, H). Sarthe: Sille le Guillaume, June 1871 (fl),
Harriot s.n. (K). Loire Inferieure: Nozay, Augusl-September 1929 (fr),
Becquet in Exsicc. Duffour 1929 801 (BM). Vendee: La Chalaigneraie, 25
July 1948 (fl), Charrier 255 (BR*, K). Loir-el-Cher: Chambord pres Blois,
1 872 (fr), Mouin s.n. (H). Cher: Acheres, 27 July 1908 (fl), Imbault s.n. (FR).
Vosges: Cremanvillers pres Vagney, Augusl 1 869 (fr), Pierrat s.n. (BM, K).
Haute Saone: St Germain, 28 Augusl 1 87 1 (fl), Vendrely s.n. (FR). Cote d' Or:
Saulieu, 27 July 1872 (fl & fr), Bonnet s.n. (K). Vienne: Monlmorillon, 19
July 1891 (fl), Violleau 669 (H). Canlal: aux environs de Quatre Chemins, 23
August 1869 (fl), Heribault-Joseph s.n. (BM). Aveyron: canton de Rignac,
montagne de Buene, 26 July 1 868 (fl), de Valon m Billet 2644 (BM). Gironde:
Biscarosse, E. of village on D652, 70 m, 8 July 1 980 (fl), M. & S. Gardner 665
(BM). Landes: Dax, lourbiere de Narosse, 23 July 1 880 (fl), Bonnet s.n. (K).
Basses Pyrenees: Biarrilz, lourbiere de Brindosse, 20 July 1880 (fl), Bonnet
s.n. (K). Hauls Pyrenees: Lac de Lourdes, June 1870 (fl), Bordere s.n. (K).
Pyrenees Orienlales: prope Biandos, Seplember 1820 (fl & fr), Bentham s.n.
(K).

ITALY. Liguria: in weslern Riviera (Gismondi, 1950: 336) bul now
exlincl (Corti, Manzi & Pedrolti, 1992). Toscano: in the Selva Pisana al San
Rossore (Corti, 1953; Pignalti, 1982: 347; Corti, Manzi & Pedrotti, 1992).

AUSTRIA. Oberoslerreich (Muhlkreis): Neuhaus a. d. Donau and Unler-
Weissenbach, perhaps extincl (Janchen, 1957: 257).

GERMANY. Bavaria: Lohrtal westlich Heigenbriicken (Spessart), 6
October 1923 (fr), Oberneder 3268 (BM). Hessen: Grossherz Hessen, bei
Obermossau im Odenwald, 9 July 1884 (fl), Durer s.n. (FR). Nordrhein-
Weslfalen: Tecklenburg, Augusl 1878 (fl), Wilmigenl s.n. (K); Gangelt
(nordlich von Aachen), 28 Augusl 1 866 (fl & fr), Becker in Wirtgen, PI. sel. fl.
Rhen. IV 175 (BM). Neidersachsen: bei Celle, Augusl 1878 (fl), Noes.n. (H);
Oldenburg, Weslerslade, June 1902 (fl & e. fr), Schiitte s.n. (H). Sachsen-
Anhall: bei Hoyerswerda, Augusl 1908 (fl & e. fr), Missbach s.n. (BM, H).

NETHERLANDS. Noord-Brabanl: Oislerwijk, N. bank of Achlerste
Choorven, 6 Augusl 1953 (fr), A"o.srer4048 (K, L). Ulrecht: between Maarn
and Maarsbergen, 16 June 1952 (fl), Kramer, Mennega & Stafleu s.n. (H).

BELGIUM. Limburg: Genek, July 1 874 (1. fl), de Dieudonne s.n. (FR,
K). Brabant: Aerschot [ Aarschol], Augusl 1 862 (fl), Devos in Van Heurck Pis
rare cril. Belg. I 1 1 (BM, K). Anlwerpen: Rijckevorsel-Canal, 12 Augusl
1920 (fr), Vermaesen s.n. (BR*, K). Namur: Canton de Gedinne, Rienne, 14
July 1861 (fl), 7 October 1861 (fr), Gravel in Req. Maill. 843 (BM, K).

CHANNEL ISLES. Jersey: Si Ouen's Pond, 11 July 1896 (fl & fr),
Lester-Garland s.n. (K). Guernsey: Grand Mare, 31 July 1914 (fl), Barton
s.n. (BM).

ENGLAND. Scilly Isles: Si Mary's, Tremelelhen (Lousley, 1971: 128).
Cornwall: Zennor, above Pendour Caslle, 2 Augusl 1947 (fl), Taylor 1 100
(K). Devon: Dartmoor above Widecombe, 9 September 1932 (e. fr), Bullard
14 (K). Dorsel: near Wareham, 5 Seplember 1957 (fl), Wycherley 222 (BM).
Hampshire: New Foresl, Brockenhurst, 15 July 1962 (fl), Whitefoord s.n.
(BM). Sussex: Ashdown Foresl, W. of Press Ridge Warren, 1 35 m, 8 Augusl
193 1 (fl), Summerhayes & Milne-Redhead 758 (K). Kenl: Chislehurst Com-
mon, 6 August 1904 (fl), Lowne 284 (K). Surrey: Ockham Common, Boulder

STUDIES IN HYPERICUM

211

Mere, 24 June 1961 (fl), Raven & Cannon 16165 (BM). Essex: Epping
Forest, August 1 883 (fl), Paulson s.n. (BM). Berkshire: Mortimer Common,
28 July 1946, Townsend s.n. (K). Suffolk: tnnear Lowestoft, 1854 (fl & fr),
Powis s.n. (BM). Norfolk: near North Walsham, August 1928 (fl & fr),
Meinertzhagen s.n. (BM). Cambridgeshire: fGamlingay, 24 August 1824
(fl), WJ. Hooker s.n. (K). Bedfordshire: Luton, 30 July 1 95 1 (fl & e. fr), Dony
s.n. (K). Gloucestershire: Ruspidge, Foxes Bridge, 20 August 1951 (fl),
Townsends.n. (K). Worcestershire: Abberley Hills, lOJune \ 840 (f\), Newman
s.n. (BM). Shropshire: Shomere, 26 July 1850 (fl), Bentham s.n. (K). Lin-
colnshire: Sawcliffe, 1857 (fr), Fowler s.n. (BM). Derbyshire: Tansley,
August 1850 (fl), Whittaker s.n. (BM, K). Cheshire: Oakmere, 8 July 1868
(fl), Bickham s.n. (BM). Lancashire: fHalsall, 4 August 1883 (fl), Brown s.n.
(LIV); tMorecambe Bay, August 1852 (fl), Lord s.n. (LIV). Yorkshire:
Skipwith Common, August 1933 (fl & fr), Riddelsdell s.n. (BM); tPilmoor,
1904 (fl & fr), Simpson s.n. (BM); tNawcliffe, 15 m, 1876 (fl), Weal s.n.
(BM). Northumberland: fBickwell Bog, August 1855 (fl), Carruthers s.n.
(BM). Westmorland: Staveley, c. 120 m, 28 July 1937 (bud), Martindale 99
(H, K). Cumberland: tDent Hill, 13 July 1868 (fl), Addison s.n. (BM).

WALES. Glamorgan: Swansea, Mayals, 25 September 1909 (fr), Riddels-
dell 289 (BM). Radnorshire: R. Wye c. 4.8 km NW of Rhyader, 1 1 August
1954 (fl), Sandwith 4155 (K). Pembrokeshire: St David's, August 1882 (fr),
Ridley s.n. (BM). Cardiganshire: Llanbadarn to Trefeglwys, 16 August 1899
(fl & fr), Marshall s.n. (BM). Montgomeryshire: Dovey Junction, 13 August
1923 (fl), Barton s.n. (BM). Merioneth: Harlech, August 1954 (fl), Boyd in
Holder 3224 (LIV). Caernarvon: Pwllheli, Llanbedrog, 6 July 1949 (fl),
Dyson s.n. (BM). Denbighshire: near Llanrust, 9 August 1930 (fl), Holder
470 (LIV). Anglesey: Holyhead I., 24 July 1853 (fl), Anon. (LIV).

ISLE OF MAN. Andreas, Gat-y-whing, June 1925 (bud), Paton s.n.
(BM).

SCOTLAND. Kirkcudbright: Palnure, Bruntis Loch, 15 August 1940
(fl), Mackechnie s.n. (K). Wigtonshire: near Port Logan, [ 1 8 — ] (fl), Balfour
s.n. (K). Ayrshire: tnear Ayr, n.d. (fl), Maclagan s.n. (BM). Stirlingshire:
tDenny Bog, 2 August 1936 (fl), Harrison s.n. (LIV). Buteshire: Isle of
Arran, near Lochranza, 25 August 1900 (1. fl), Marshall s.n. (BM). Argyll-
shire: Isle of Gigha, 1898 (fr), Somerville s.n. (BM); Isle of Islay, east side, 5
September 1 883 (fl), Ley s.n. (BM); Isle of Mull, Ross of Mull, CamasTuath,
7 July 1968 (fl), BM Mull Survey 2904 (BM). Invernessshire: Isle of Canna,
August 1897 (fr), Macvicar s.n. (BM). Outer Hebrides: Harris, W. coast, 8-
1 5 August 1959JutePerring mProc. B. S.B.I. 4: 206 ( 1961); N. Uist, Newton,
November 1 936 (o. fr), Campbell s.n. (BM); S. Uist, 1 936 (e. fr), Comber s.n.
(BM); Barra, 5 July 1887 (fl), Somerville s.n. (BM); Vatersay, near sea, 17
August 1983 (fl & fr), J. & M. Cannon & Charley 2 (BM).

IRELAND. Kerry: near Lough Guitane, c. 90 m, 5 September 1935 (fl &
fr), Ross-Craig, Burn & Sealy 374 (K); Caragh, 4 July 1970 (fl), Greenwood
s.n. (LIV). Cork: near Glengariff, August 1935 (fl & fr), Ross s.n. (BM).
Waterford: Carrickovantry, c. 3.2 km W. of Tramore, 27 August 1967 (fl),
Ferguson 2070 (BM). Tipperary: fClogheen, Castlegraco, August 1850 (fl),
Carroll s.n. (BM). Galway : Connemara, Rounstone, August 1 884 (fl), Green
s.n. (LIV). Wicklow: Croghan Mt., July 1882 (fl), Fawcett s.n. (BM). Louth:
tCarlingford, 22 August 1895 (fl), Last s.n. (K). Donegal: tKilmacrenan,
August 1 886 (fl), Kinshan s.n. (BM). Down: fNewtonards, August 1 87 1 (fl),
Linton s.n. (BM).

AZORES. Sao Miguel: Lagoa dos Furnas, 400 m, 15 July 1972 (fl),
Gonfalves 4318 (BM); Lagoa do Junco, 770 m, 30 July 1970 (fl), Dolman
386 (BM). Pico: Pico, c. 600 m, 1 September 1970 (1. fl), Brooke 1 1 323 (BM).

H. elodes is clearly derived from sect. Adenosepalum, its nearest
relative being 13. H. coadunatum, from the Canary Islands. With
regard to vegetative characters, H. elodes is more advanced than the
H. caprifolium group (Spp. 13-15), and its occurrence in wet
habitats is also a more extreme expression of a tendency shared with
that group. The pseudo-tubular flower structure of H. elodes, how-
ever, is quite distinct, although Webb & Berthelot's figure of H.
coadunatum suggests that the flower of that species may sometimes
be basally contracted. The possession of a full-fledged syndrome of
characters associated with specialized pollination, however, distin-

11 1 Extinct.

guishes H. elodes from all other species of Hypericum except those
in the distantly related sect. 25. Adenotrias (Robson, 1972, 1981:
1 22) and justifies its position in an admittedly paraphyletic monotypic
section. Two distinct basic chromosome numbers have been re-
corded for this species (n = 10, 8), see p. 209.

The Greek 'eXo8r|^ has been transliterated by various authors as
elodes or helodes, the Greek rough breathing (spiritus asper) sign '
thus being replaced by h, which it represents. Although this practice
is to be recommended when coining new names based on Greek, an
author's original spelling should be retained, and Linnaeus (follow-
ing Clusius) used the form elodes. St-Lager ( 1 880) and Mansfeld
(1939) were therefore wrong to correct it to helodes.

The habitats of H. elodes are everywhere being increasingly
threatened by drainage and water-extraction, and so its distributional
area has been considerably reduced in the last century or so. The
changes are most noticeable in the drier east (e.g. in Germany and
north-east England), but they are occurring throughout the range of
the species.

Gliick (1911) described three forms of H. elodes that are related
to the depth of water in which it is growing. The shallow-water
form is finely pubescent with a long, creeping stem bearing ob-
long-elliptic leaves and giving rise to emergent erect shoots. These
bear broadly ovate, often cordate leaves and terminal (though often
apparently axillary) inflorescences. Where the water has dried up
or the vegetation is dense, the shallow-water form is replaced by
the terrestrial form (later named H. helodes forma terrestre Gliick),
which is essentially similar but smaller and more reduced. In both
these forms the creeping stems with 'primary' leaves can
overwinter, producing aerial shoots the following summer. Where
the water is deeper (up to c. 50 cm), a deep-water form occurs
(Elodes palustris forma submersa Gliick). In it the water shoots,
which can exceed 40 cm in length, are paired, unbranched,
filamentous, glabrous and sterile, with leaves narrowly to broadly
elliptic. Proper leaves are mostly confined to the upper half of the
shoot, those toward the base being small or even almost scale-like.
If these shoots reach the surface of the water, they can immediately
become aerial shoots. Hagemann (1983: 130) likewise demon-
strated that the original shoots are plagiotropic, and that lateral
shoots from them both (i) terminate in an inflorescence and (ii)
produce axillary shoots that flower in the second year.

The seeds of//, elodes are shed in August (in Germany), float for
about three days, and then sink and overwinter in the mud at the
bottom of the pond, etc. They germinate early the following year and
are not inhibited by having dried out in the meantime.

ACKNOWLEDGEMENTS. My studies for Part 6 have been especially helped
by Dr W. Preston Adams, whose work on sect. Myriandra (including his
unpublished Ph.D thesis) has been the basis of my own on that section, and Dr
Peter Raven (MO), who has facilitated my work on North American species
in several ways. Others whose assistance in my studies I should like to
acknowledge in particular are: Mrs Sheila Collenette (Arabia), the late Dr
Peter Davis (E), Dr Isolde Hagemann (B), Dr Richard Howard (A), Dr
Stephen Jury (RNG), the late Dr Bassett Maguire (NY), DrTony Miller (E),
Dr A. Ramos Nunez (MA), Mme Claude Reynaud (MARS), Dr Friederike
Sorger (Vienna, collection now at W) and Dr Peter Stevens (GH), as well as
my colleagues Prof. Chris Humphries, Drs Mike Gilbert, Rob Huxley and
Charlie Jarvis and also Nick Turland and Peter Stafford. My thanks are also
due to the directors of the following herbaria for loans and (to some) for study
facilities: A, BISH, BO, CANB, CM, DUKE, E, F, FR, G, GH, H, JE, K, L,
LISJC, MARS, MEL, MEXU, MICH, MO, NY, P, PNH, PRE, RNG, S,
SING, RGH, UPS, US, W, WIS, WRSL and Z, and to the late Dr A. Huber-
Morath (Basel, collection now at BASBG). I am again deeply indebted to Mrs
Margaret Tebbs for drawing the plates and figures, and I am also very grateful

212

N.K.B. ROBSON

to Ms Marian Short, Mrs Jenny Gunn and Dr Richard Bucknall for help with
word-processing, and Miss Siri France for assistance with the text illustra-
tions. My wife Eve has, as always, been a help and support in many ways. Last
but not least, I must thank the Keeper of Botany, Dr Stephen Blackmore, for
study facilities in the Botany Department, and the Linnean Society for a grant
from the Appleyard Fund.

REFERENCES

Adams, W.P. 1957. A revision of the genus Axcyrum (Hypericaceae). Rhodora 59: 73-
95.

1959. The taxonomy of Hypericum section Myriandra (Hypericaceae). Unpub-
lished thesis. Harvard University.

1962. Studies in the Guttiferae. I. A synopsis of Hypericum section Myriandra.

Contr. Gray Herb. Han: no. 189: 1-51.

1972. Studies in the Guttiferae. III. An evaluation of some putative spontaneous

garden hybrids in Hypericum section Myriandra. Rhodora 74: 276-282.

— 1973. Clusiaceae of the Southeastern United States./ Elisha Mitchell sci. Soc. 89:

62-71.
& Robson, N.K.B. 1961. A re-evaluation of the generic status of Ascyrum and

Crookea (Guttiferae). Rhodora 63: 10-16.
Al-Bermani, A.-K.K.A., Al-Shammary, K.I.A., Bailey, J.P. & Gornall, R.J. 1993.

Contributions to a cytological catalogue of the British and Irish flora, 3. Watsonia 19:

267-278.
Arrigoni, P.V. 1965. Richerche geobotaniche su Linaria miilleri Moris e notizie su

Hypericum aegyptiacum L., nuovo reperto per la flora sarda. Webbia 20: 307-330.
Corrias, B., Corrias, S.D., Nardi, E. & Valsecchi, F. 1973. Nuovo stazioni di

Hypericum annulatum Moris e Ribes multiflorum Kit. ssp. sandalioticum Arrig. in

Sardegna. Webbia 28: 423^25.
Audley-Charles, M.G. 1987. Dispersal of Gondwanaland: relevance to evolution of

the angiosperms. In T.C. Whitmore (Ed.), Biogeographical evolution of the Malayan

Archipelago: 5-25. Oxford.
Balfour, I.B. 1888. Botany of Socotra. Trans. R. Soc. Edinb. 31. Pp. Ixxvi + 446 & tt.

100.
Bamps, P. 1970. Guttiferae. In R. Boutique (Ed.), Flare du Congo du Rwanda et du

Burundi. Spermatophytes. Bruxelles.
Robson, N.K.B. & Verdcourt, B. 1978. Guttiferae. In R.M. Polhill (Ed.), Flora

of Tropical East Africa. London.
Barker, C. & Cheek, M. 1994. Plant portraits: 245. Hypericum buckleyi. Guttiferae.

Ken-Mag. 11:65-69,1.245.

Boissier, E. 1867. Flora Orientalis 1, Thalamiflorae. Basel.
Bonafe Barcelo, F. 1979. Flora Mallorca 3. Palma de Mallorca.
Borg, J. 1927. Flora of the Maltese Islands. Malta.
Borgen, L. 1969. Chromosome numbers of vascular plants from the Canary Islands,

with special reference to the occurrence of polyploidy. Nytt Mag. Bot. 16: 81-121.
Borgmann, E. 1964. Anteil der Polyploiden in der Flora des Bismarckgebirges von

Ostneuguinea. Z. Bot. 52: 1 18-172.

Bostick, P.E. 1965. Documented plant chromosome numbers, 65: 2. Sida 2: 165-168.
Bredell, H.C. 1 939. A revision of the South African species of Hypericum. Bothalia 3:

571-182.

Briquet, J. 1935. Prodrome de la Flore Corse, Guttiferae. 2(2): 143-154.
Britton, N.L. 1918. Flora of Bermuda. New York.
& Brown, A. 1913. An illustrated Flora of the northern United States, Canada

and the British possessions (etc.) 2 (Amaranthaceae to Loganiaceae). New York.
Burdet, H.ML, Charpin, A. & Jacquemoud, F. 1984. Types nomenclaturaux de taxa

iberiques decrits par Boissier ou Reuter. VI. Euphorbiacees a Guttiteres. Candollea

39: 771-789.

Clark, R.C. 1971. The woody plants of Alabama. Ann. Mo. hot. Gdn 58: 99-242.
Cooperrider, T.S. 1989. The Clusiaceae of Ohio. Castanea 54: 1-11.
Cornell, D.S. & Johnston, M.C. 1 970. Manual of the vascular plants of Texas. Renner.
Corti, R. 1953. Su H\pericum elodes Huds., relitto eu-atlantico della selva di San

Rossore (Pisa). AttiAccad. naz. Lincei, Ser. 8, Rend. cl. sci., mat. e nat. 14: 308-314

with map & 3 tt.

Corti, R., Manzi A. & Pedrotti, F. 1992. Libm rosso delle piante d' Italia. Roma.
Coulter, J.M. 1886a. Revision of North American Hypericaceae. I. Bot. Gaz. 11: 78-

88.

18866. Some notes on Hypericum. Bot. Gaz. 11: 275.

Dalgaard, V. 1986. Chromosome studies in flowering plants from Madeira. Willdenowia

16: 221-240.
1 99 1 . Chromosome studies in flowering plants from Macaronesia. II . Willdenowia

20: 139-152.
Delay, J. 1972. Prospection caryologique en Brenne et Limousin. Bull. soc. hot. France

116 (97C sess. extraord.): 69-74.
Fawcett, W. & Rendle, A.B. 1926. Flora of Jamaica 5. London.

Fernald, M.L. 1950. Gray's manual of botany 8th ed. New York.

& Schubert, B.C. 1948. Studies of American types in British herbaria. III.

Rhodora 50: 181-208.

Forbes, H.O. 1903. The natural history of Socotra and Abd-el-kuri. Liverpool.

Gagnieu, A. & Wilhelm, J.P. 1965. Genre Hypericum [In A. Gagnieu, Les chromo-
somes dans la cellule, la piante, 1'espece], In Anon. (Ed.), Travaux biologiques dedit
a Prof. Plantefol: 472-473. Paris.

Galland, N. 1988. Recherche sur 1'origine de la flore orophile du Maroc; etude
caryologique et cytologique. Trav. Inst. Sci. Univ. Mohammed V, Bot. (Rabat) 35: I-
168.

Gamisans, J. 1985. Catalogue des plantes vasculaires de la Corse. Ajaccio.

Gattinger, A. 1887. The Tennessee Flora. Nashville.

Gillespie, J.P. 1958. The Hypericaceae of Tennessee. Castanea 24: 24-32.

Gillett, J.M. 1975,/n A. Love (Ed.), I.O.P.B. chromosome number reports, L. Taxon 24:
671-678.

— & Robson, N.K.B. 1981. The St. John's-worts of Canada (Guttiferae). Canad.
natn. Mus. nat. Sci., Publs Bot., No. 1 1. Ottawa.

Gimingham, C.H. & Walton, K. 1954. Environment and the structure of scrub

communities on the limestone plateaux of northern Cyrenaica. J. Ecol. 42: 505-520.
Gismondi, A. 1950. P rospetto della Flora Ligustica. Genoa.
Gliick, H. 191 1. Biologische und morphologische Untersuchungen iiber Wasser- und

Sumpfgewachse, 3. Die Uferflora. Jena.
Gorschkova, S.G. 1949. Guttiferae. In B.K. Shishkin & E.G. Bobrov (Eds), Flora S. S.

S. R. 15: 201-258.
Greuter,W.,Matthas,W.& Risse,H. 1984. Additions to the flora of Crete, 1973-1983

(1984) - II. Willdenowia 14: 269-297.

— Pfleger, R. & Raus, T. 1983. The vascular flora of the Karpathos island group
(Dodecanesos, Greece). A preliminary checklist. Willdenowia 13: 43-78.

Grossheim, A.A. 1962. Hypericaceae. Flora Kavkasa 2nd ed., 6: 163-178. Baku.
Hagemann, I. 1983. Wuchsformenuntersuchungen an zentraleuropaischen Hyperi-
cum- Arten. Flora 173: 97-142.
1 989. Wuchsformen einiger Hypericum- Arten, ein Beitrag zum morphologischen

und zum okologischen Anliegen der Wuchsformen-Forschung. Flora 183: 225-309.
Haslam, S., Sell, P.D. & Wolseley, P.A. 1977. A flora of the Maltese Islands. Maida.
Hedberg, I. & O. 1977. Chromosome numbers of afroalpine and afromontane

angiosperms. Bot. Notiser 130: 1-24.
Hiepko, P. 1979. Das Schiksal des 'Ausserrheinischen Herbariums' des

Naturhistorischen Vereins fur die Preussischen Rheinlande und Westfalen.

Willdenowia 9: 207-208.
Hoar, C.S. & Haertl, E.J. 1932. Meiosis in the genus Hypericum. Bot. Gaz. 93: 197-

205.

Hooker, J.D. 1 880. Hypericum aegyptiacum. Curtis Bot. Mag. 106: t. 648 1 .
Jacobs, M. 1 972. The plant world on Luzon 's highest mountains. Leiden.
Jacot Guillardmot, A. 1971. Flora of Lesotho (Basutoland). Lehre.
Janchen, E. 1957. Hypericum. In F. Knoll & K. Hofler, Catalogus Florae Austriae 1:

256-257.
Jaubert, H.-F., Comte de & Spach, E. 1842-1843. Illust rat tones plantarum

Orientalium 1. Paris.
Jennings, O.E. 1917. II. A contribution to the botany of the Isle of Pines, Cuba [etc.].

Ann. Carneg. Mus. 11: 19-290.

Kask, M. 1971. Guttiferales. In K. Eichwald et al. (Eds), Eesti NSV Floora 8: 19-35.
Keller, R. 1 893. Hypericum. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien

3(6): 208-215.
1 925. Hypericum. In A. Engler & K. Prantl, Die naturlichen Pflanzenfamilien 2nd

ed. 21: 175-183.
Killick, D.J.B. & Robson, N.K.B. 1976. Clusiaceae. In J.H. Ross (Ed.), Flora of

Southern Africa 22: 14-24.
Kimura. Y. 1951. No. 10, Hypericaceae. In T. Nakai & M. Honda (Eds), Nova flora

japonica. Tokyo.

Kunkel, G. 1977. Endemismos Canarios. Madrid.
Larsen, K. 1962. Contribution to the cytology of the endemic Canarian element. Bot.

Notiser US: 196-202.
Lewis, W.H., Stripling, H.S. & Ross, R.G. 1962. Chromosome numbers of some

angiosperms of the southern United States and Mexico. Rhodora 64: 147-161.
Li Xi-wen 1990. Guttiferae. In Li Xi-wen (Ed.), Flora Reipublicae Popularis Sinicae

50(2): 1-112.
Lippold, H. 1970. Die Hypericum-Arten Cubas. Wiss. z. Friedrich-Schiller-Univ. Jena,

Math.-Nat. R. 19: 377-382.
Loon, J.C. van & de Jong, H. 1978. In A. Love (Ed.), I.O.P.B. chromosome number

reports, LIX. Taxon 27: 53-61.

Lousley, J.E. 1971. The Flora of the Isles of Sc illy. Newton Abbott.
Love, A. & D. 1982. In A. Love (Ed.), I.O.P.B. chromosome number reports, LXXV.

Taxon 31: 342-368.

— & Kjellqvist, E. 1974. Cytotaxonomy of Spanish plants. IV. Dicotyledons:

Caesalpiniaceae - Asteraceae. Lagascalia 4: 153-21 1.
Maire, R. 1924. Etudes sur la vegetation et la flore du Grande Atlas et du Moyen Atlas

marocains. Mem. Soc. Sci. nat. Maroc no. 7: 1-220 & tt. 16. Rabat.

STUDIES IN HYPERICUM

213

1933. Etudes sur la flore et la vegetation du Sahara central. Mem. Sot: Hist. mil.

Afr. Nurd no. 3: 154.
Mansfeld, R. 1939. Zur Nomenklatur der Farn- und Blutenpflanzen Deutschlands.

Feddes Reprium Spec: nov. veg. 47: 263-287.
Melville, R. 1967. The distribution of land around the Tethys Sea and its bearing on

modern plant distribution. In C.G. Adams & D.V. Ager (Eds), Aspects of Tethvan

biogeography. 291-312. Svst. Ass. Publ. no. 7. London.
Mennema, J., Quene-Broterenbrood, A.J. & Plate, C.L. 1980. (Eds), Atlas of the

Netherlands flora 1. The Hague.
Millspaugh, C. F. 1892. Preliminary catalogue of the flora of West Virginia. Bull. W. Va.

Exp.Sta. 24:311-537.
Milne-Redhead, E. 1953. Hypericaceae. In W.B. Turrill & E. Milne-Redhead (Eds),

Flora of Tropical East Africa. London.
Moggi, G. & Pisacchi, A. 1967. Adumbratio Florae Aethiopicae. 14. Hypericaceae.

Webbia 22: 233-289.
Mohlenbrock, R.H. & Evans, O.K. 1972. Illinois field and herbarium studies.

Rhodora 74: 142-151.

& Voigt, J.W. 1959. A Flora of Southern Illinois. Carbondale.

Morley, RJ. & Flenley, J.R. 1987. Late Cainozoic vegetational changes in the

Malayan Archipelago. In T.C. Whitmore (Ed.), Biogeographical evolution of the

Malav Archipelago: 50-59. Oxford.

Mouterde, P. 1970. Nouvelle Flore du Liban et de la Syrie, 2(texte). Beirut.
Munz, P.A. 1974. A Flora of Southern California. Berkeley.
Myers, O., Jr. 1963. Studies of reproduction and hybridization in five species of

Hypericum native to Eastern North America. Ph.D. Thesis, Cornell Univ. (See Diss.

A tor. 24:3515-3516, 1963).
Nielsen, N. 1924. Chromosome numbers in the genus Hypericum. (A preliminary

note.) Hereditas 5: 378-382.

Noack, K.L. 1934. Uber Hypericum-Kremungen. IV. Die Bastarde zwischen Hyperi-
cum acutum Monch, montanum L., quadrangulum L., hirsutum L. und pulchrum L.

Z. Bot. 28: 1-71.
1939. Uber//y/w;'tMW-Kreuzungen. VI. Fortpflanzungsverhaltnisse und Bastarde

von H. perforatum L. Z. indukt. Abstamm.- u. VererbLehre 76: 569-601.
Ohlendorf, O. 1907. Beit rage zur Anatomie und Biologic der Fruchte und Samen

einheimischer Wasser- und Sumpfpflanzen. Inaug. Diss. Univ. Erlangen.
Ortega, J. & Navarro, B. 1978. Estudios en la flora de Macaronesia: algunos numeros

de cromosomas. III. Botanica macaron. 3: 73-80.

Pignatti S. & Moggi, G. 1982. In S. Pignatti, Flora d' Italia, 1: 343-351. Bologna.
Pettier- Alapetite, G. 1979. Flore de la Tunisie. 1. Angiosperms - Dicotyledones,

Apetales - Dialypetales. Tunis.
Pringle, J.S. 1976. Annotated chromosome counts for some plants of the dunes and

pannes along the shores of the upper Great Lakes. Michigan Botanist 15: 157-163.
Queiros, M. 1991. Numeros cromosomicos de algumas especies Portuguesas de

Hypericum. Revta Biol. Univ. Oviedo 9: 51-57.
Quezel, P. & Santa, S. 1963. Nouvelle flore de I'Algerie 2. Paris.
Radford, A.E., Ahles, H.E. & Bell, C.R. 1968. Manual of the vascular flora of the

Carolinas. Chapel Hill.

Rafinesque-Schmaltz, C.S. 1817. Flora Ludoviciana. New York.
Ramos Nunez, A.F. 1983 ['1982']. Estudio biosystematico del genero Hypericum L.

(Guttiferae) en la Peninsula Iberica e Islas Baleares. I. Caracteres seminales. Trab.

Dep. Bot. Univ. Complut. 12: 45-62.
1987. Clusiaceae. In B. Valdes, S. Talavera & E.F. Galiano. (Eds), Flora vascular

del Andalucfa occidental 1: 314—318.

1993. Hypericum. In S. Castroviejo et al. (Eds), Flora Iberica 3: 157-185.

Madrid.

Rehder, A. 1911. Pistillody of stamens in Hypericum nudiflorum. Bot. Gaz. 51: 230-

231.
Reynaud, C. 1973. Contribution a 1'etude cytotaxinomique du genre Hypericum L. en

Turquie. I. Bull. Soc. hot. France 120: 201-216.
1975. Contribution a 1'etude taxinomique de quelques//v/>en'cw/fl mediterraneans.

Biol.-Ecol. medit. 2: 3-8.
1980. Contribution a 1'etude cytotaxinomique du genre Hypericum L. en Grece.

Bull. Soc. hot. France 127: 345-353.
198 1 . Contribution a 1'etude cytotaxinomique du genre Hypericum en Turquie. II.

Biol.-Ecol. medit. 8: 181-191.

1985. Etude des teguments seminaux de quelques Hypericum (Guttiferae)

mediterraneens observes au M.E.B. I. Bull. Mus. natn. Hist, not., Paris IV, 7 sect. B,
Adansonia no. 1: 85-96.

1986. Etude cytotaxinomique des Millepertuis du Bassin mediterraneen et des

lies Canaries. Bull. Soc. hot. France 133, Lettres Bot. 1986 (2): 167-177.

1991. Etude des teguments seminaux (observes au M.E.B.) de quelques Hyperi-
cum (Guttiferae) mediterraneens. II. Bull. Mus. natn. Hist, not., Paris IV, 13 sect. B.,
Adansonia no. 3^: 183-195.

Robin, C.C. 1807. Voyage dans I'interieurde la Louisiana[. . .] 3. Flore louisianaise.
Paris.

Robson, N.K.B. 1956. Studies in the genus Hypericum L Unpublished thesis, Edin-
burgh Univ.

1 958. The genus Hypericum in Africa south of the Sahara, Madagascar and the

Mascarenes. Kew Bull. 12: 443-446.

1967«. Materials for a flora of Turkey: XI. Notes on Turkish species of Hyperi-
cum. Notes R. hot. Gdn Edinb. 27: 185-204.

I967fe Hypericum. In P.H. Davis (Ed.), Flora of Turkey and the east Aegean

Islands 2: 355-401. Edinburgh.

— 1968. Guttiferae (Clusiaceae). In T. Tutin et al. (Eds), Flora Europaea 2: 261-
269.

1972. Evolutionary recall in Hvpericum (Guttiferae)? Trans, hot. Soc. Edinb. 41:

365-383.

1973 ['1972']. Notes on Malesian species of Hypericum (Guttiferae). Rora

Malesiana Praecursores, LII. Blumea 20: 251-274.

— 1974. Hypericaceae. In C.G.G.J. van Steenis (Ed.), Flora Malesiana I, 8: 1-29.
1977a. Studies in the genus Hypericum L. (Guttiferae). 1 . Infrageneric classifica-
tion. Bull. Br. Mus. nat. Hist. (Bot.) 5: 291-355.

1977fe Notes on some Nepalese and Indian Hypericum. J. Jap. Bot. 52: 276-288.

1980. The Linnaean species of Ascyrum (Guttiferae). Taxon 29: 267-274.

1 98 1 . Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the genus.

Bull. Br. Mus. nat. Hist. (Bot.) 8: 55-226.
1 985. Studies in the genusHypericum L. (Guttiferae). 3. Sections 1 . Campylosporus

to 6a. Umbraculoides. Bull. Br. Mus. nat. Hist. (Bot.) 12: 163-325.
1987. Studies in the genus Hypericum L. (Guttiferae). 7. Section 29. Brathys (part

1). Bull. Br. Mus. nat. Hist. (Bot.) 16: 1-106.

1990. Studies in the genus Hypericum L. (Guttiferae). 8. Sections 29. Brathys

(part 2) and 30. Trigynobrathys. Bull. Br. Mus. nat. Hist. (Bot.) 20: 1-151.

1992. Hypericum. In A.H. Huxley (Ed.), The New Royal Horticultural Society

Dictionary of Gardening 2: 620-637.

1993a. Parallel evolution in tropical montane Hvpericum. Opera Botanica 121:

263-274.

19936. Studies in Hypericum: validation of new names. Bull. nat. Hist. Mus. Lond.

(Bot.) 23: 67-70.

1995. Hypericum. In J. Cullen et al. (Eds), European Garden Flora 4: 44-74.

& Adams, W.P. 1968. Chromosome numbers in Hypericum and related genera.

Brittonia 20: 95-106.
Rompaey, E. van & Delvosalle, L. 1979. Atlas de la flore beige et luxumbourgois,

Pteridophytes et Spermatoph\tes. 2nd ed. Meise.
St.-Lager, J.B. 1880. Retorme de la nomenclature botanique. Annls Soc. hot. Lyon 7:

1-154.
Saunders, E.R. 1937. The non-unique position in the genus Hypericum of H. peplidi-

folium A. Rich. Proc. Linn. Soc. London 149: 160-163.
Small, J.K. 1933. Manual of the southeastern Flora. New York.
Smith, A.C. 1941. Studies in Papuasian plants, III. Guttiferae. J. Arnold Arbor. 22:

343-374.
Smith, J.M.B. 1 977. Origins and ecology of the tropicalpine flora of Mt. Wilhelm, New

Guinea. Biol. J. Linn. Soc. 9: 87-131.
1986. Origins and history of the Malesian high mountain flora. In F. Vuilleumier

& M. Monasterio (Eds), High altitude tropical biogeography: 469—477. Oxford.
Soo, R. 1 968. Hypericum in Synopsis Systematico-Geobotanico Florae Vegetationisque

Hungariae 3: 431-438.
Spach, E. 1836fl. Histoire naturelle des vegetaux. Phanerogames 5: 335^64.

1836b. Conspectus monographiae Hypericacearum. Annls Sci. nat. (Bot.) II, 5:

349_369, t. 6.

Steenis, C.J J.G. van 1 964. Plant geography of the mountain flora of Mt. Kinabalu.

Proc. Roy. Soc. London B, 161: 7-38.
StefanofT, B. 1931. Notes on new or rare Hypericum. Kew Bull. 1931: 29-33.

1932-1934. Sistematicheski i geografiski prouchvaniya verchu mediterransko-

orientaliskite vidovye na roda Hypericum L. God. Agr.-les. Fak. Univ. Sofiya 10:
19-57 (1932); 11: 130-186 (1933a); 12: 69-100 (1934).

1933b. Die mediterran-orientalischen Arten der Gattung Hypericum L.

Pflanzenareale 4: Karten 1-9.

Stjepanovi6Vesilichi6, L. 1972. Hypericaceae. In M. Josifovi6 (Ed.), Flora S. R..

Srbije3: 102-125.
Stoker, F. 1 95 1 . In F.J . Chittenden (Ed. ), The Royal Horticultural Society Dictionary of

Gardening 2: 1033-1038.
Strid, A. & Kraiiy.cn, R. 198 1 . In A. Love (Ed.), I.O.P.B. chromosome number reports,

LXXIII. Taxon 30: 829-842.
Svenson, H.K. 1940. Plants of the southern United States. III. Woody species of

Hypericum. Rhodora 42: 8-19.

1952. What is Hypericum prolificum! Rhodora 54: 205-207.

Torrey, J. & Gray, A. 1838. Hypericaceae in Flora of North America 1: 155-169;

1840, Supplement: 671-674.
Turland, N. 1990. Cretan Bellflowers and St. John's Worts. Quart. Bull. A. G. S. 58:

310-320.

1992. Floristic notes from Crete. Bot. J. Linn. Soc. 108: 345-357.

Utech, F.H. & Iltis, H.H. 1970. Preliminary reports on the flora of Wisconsin, No. 61.

Hypericaceae - St. John's Wort family. Trans. Wis. Acad. Sci. Arts Lett. 58: 325-35 1 .
Voggenreiter, V. 1974. Geobotanische Untersuchungen an der naturlichen Vegetation

214

N.K.B. ROBSON

der Kanareninsel Tenerit'e [. . .] als Grundlage fur den Naturschutz. Dissert, hot. 26.
Vulf, E.V. 1953. Guttiferae in Flora Kryma 2(3): 106-1 1 1.
Ward, D.B. 1980. In D.B. Ward (Ed.), Rare and endangered biota of Florida 5: Plants.

Gainesville.

Webb, P.B. & Berthelot, S. 1836. Histoire naturelle dex lies Canaries 3(2).

Phytographia canarienxis, 1 . Paris.
Wilbur, R.L. 1995. The orthography of the name of a southeastern endemic shrub

Hypericum buckleii M.A. Curtis. Castanea 60: 168-169.

SYSTEMATIC INDEX

Accepted names are in roman and synonyms in italic; new names and principal references are in bold. An asterisk (*) denotes a figure. Adm = sect.
Adenosepalum, Ads = sect. Adenotrias, Ar = sect. Arthrophyllum, E = sect. Elodes, He = sect. Heterophylla, Hu - sect. Humifusoideum, M = sect. Myriandra,
T = sect. Triadenioides, W = sect. Webbia.

Adenosepalum montanum (L.) Fourr. (=Adml7) 196

tomento.sum (L.) Fourr. (=Adm9) 186
Adenotrias kotschvi Jaub. & Spach (=Ads2) 152

phrygia Jaub. & Spach (=Ads2) 152
Ascyrum group (M25-29) 78, 118
Axcvrum L. (=sect. 20 subsect. 5)

a. Ascyrum (L.) Endl. (=sect. 20 subsect. 5) 124

b. Ixophvllum (Spach) Endl. (=sect. 20 subsect.

3) 113
amplexicaule Michx. (=M26) 127

sensu Pursh, pro pane (=M25) 125
cruciatum St-Lag. (=M25) 125
crux-andreae L. (=M25) 124

sensu Chapm. (=M29a) 131

sensu Coulter (=M29b) 132

[var.] (3 angustifulium Null. (=M29a) 131

[var.] (3 sensu Choisy, pro parte (=M29b) 132
cubense Griseb. (=M26) 127
cuneifolium Chapm. (=M25) 125, 127, 129
edisonianum Small (=M27) 128
filicaule Dyer 124
grandiflorum Raf. (=M25) 125
helianthemifolium Spach (=M29b) 132
hypericoides L. (=M29) 124, 129

sensu W.T. Aiton (=M25) 125

var. multicaule (Michx. ex Willd.) Fernald
(=M29b) 132

var. oblongifolium (Spach) Fernald (=M29a) 131

var. rypicum Fernald (=M29a) 131
linifolium Spach (=M29a) 131, 132
macmsepalum S. Brown (=M29a) 131, 132
michauxii Spach (=M29a) 131
microsepalum Torrey & Gray (=M19) 1 17
montanum Raf. (=M29a) 131
multicaule Michx. ex Willd. (=M29b) 132
nummularifolium Banks ex Steud. (=M28) 129
oblongifolium Spach (=M29a) 131
pauciflorum Nutt. (=M28) 129
perforata Lam. (=M29) 130
plumieri Bertol. (=29a) 131
pumilum Michx. (=M28) 129
simplex Zeyh. ex Turcz. (=M25) 125
xpathulatum Spach (=M29b) 132
stans Michx. ex Willd. (=M25) 124, 127

[var.] p obovatum Chapm. ex Torrey & Gray
(=M25) 125, 127

[var.] p sensu Choisy (=M26) 127
tetrapetalum (Lam.) Vail (=M26) 127
villosum L. 124
Ascyrum foliis lanceolatis-linearibus (etc.) Burm.

(=M29c) 133

Ascyrum foliis oblongis (etc.) Burm. (=M29a) 131
BetulaL. 110

Brathydium Spach (=sect. 20 subsect. 4) 122
amhiguum (Elliott) K. Koch (=M6) 102
aureum (W. Bartram) K. Koch (=M1) 94
canadense Spach ( =M22 ) 121
chamaenerium Spach (=M20) 1 1 8
chamaerinum sensu Steud. 1 1 8
dalabriforme (Spach) Y. Kimura (=M24) 123
fastigiatum (Elliott) K. Koch (=M21) 120
fulgidum (Raf.) K. Koch (=M13?) 1 1 1
grandiflorum Spach (=M24) 122, 123

hyssop/folium Spach (=M18) 116
microsepalum (Torrey & Gray) K. Koch

(=M19) 117

myrtifolium (Lam.) K. Koch (=M23) 122
nudiflorum (Michx. ex Willd.) K. Koch

(=M17) 114

rugelianum (Kun/,e) K. Koch (=M1) 94
sphaerocarpum (Michx. ex Willd.) Spach

(=M20) 118

Brathyx prolifica (L.) Payer (=M2) 97
Cistus laurifolius L. 147
Cmokea Small (=sect. 20 subsect. 3) 1 13

microsepala (Torrey & Gray) Small (=M19) 1 13,

117

Croton elaeagnoides Balf. f. 143
Elodea aegyptica (L.) Jack (=Adsl) 149
palustris i. St. Hill. (=E1) 209
russeggeri (Fenzl) Walp. (=Ads2) 152
Elodes acifera Greuter (=Ads3) 153
aegyptica (L.) Payer (=Adsl) 149
palustris Spach (=E1) 209

forma submersa Cluck (=E1 ) 209, 21 1
russeggeri (Fenzl) Greuter (=Ads2) 152
Episiphis parvifolia Raf. (=Adsl) 149
Helodes glandulosum St.-Lager (=E1) 209
H\pericoides Adans. (=sect. 20 subsect. 5) 124
crux-andreae (L.) Poir. (M=25) 125
perforata Poir. (=M29) 124,130
Hypericoides frutescens, erecta (etc.) Plum.

(=M29a) 131
Hypericoides frutescens, humi-fusa (etc.) Plum.

(=M29c) 133
Hypericum L. 75, 76, 77, 78, 88, 118, 147, 179

sect. 27. Adenosepalum Spach 76, 88, 89, 90, 170,
175, 179, 199,211
subsect. Adenosepalum 90, 193
subsect. Aethiopica N. Robson 90, 172
subsect. Caprifolia N. Robson 90, 189
subsect. Pubescentes N. Robson 90, 181
sect. 25. Adenotrias (Jaub. & Spach) R. Keller 76,

83,84,85, 147, 150, 153,211
sect. 22. Arthrophyllum Jaub. & Spach 76, 82, 83,

90, 137, 138, 140
sect. 3. Ascyreia Choisy 76
sect. Brathydium (Spach) R. Keller (=sect. 20) 77
subsect. Eubrathydium R. Keller (=sect. 20

subsect. 4) 77
subsect. Pseudobrathydium R. Keller (=sect. 20

subsect. 2) 77

sect. 29. Brathys (Mutis ex L. f.) Choisy 133
sect. 8. Bupleuroides Stefanoff 76
sect. 1. Campylosporus (Spach) R. Keller 76, 78,

79,87,90, 135, 145, 146, 157, 175
sect. 13. Drosocarpium Spach 198
sect. 28. Elodes (Adans.) W. Koch 76, 88, 89, 90,

208
sect. Euhvpericum Boiss. 76

subsect. Homotaenium R. Keller 76
sect. 24. Heterophylla N. Robson 76, 83, 84, 88,

146

sect. 17. Hirtella Stefanoff 76
sect. 26. Humifusoideum R. Keller 76, 85, 86, 87,
88, 153, 154, 157, 159, 162

sect. 9. Hypericum 76, 90, 154, 179

sensu N. Robson, pro parte (=sect. 26) 153
sect. Isophyllum (Spach) W.P. Adams (=sect. 20

subsect. 3) 113
sect. 20. Myriandra (Spach) R. Keller 76, 77, 78,

79,92

subsect. 5. Ascyrum (L.) N. Robson 78, 80, 124

subsect. 4. Brathydium (Spach) R. Keller 78, 80,
82, 122

subsect. 1. Centrosperma R. Keller 78, 80, 81,
94,98, 123

subsect. Centrosperma sensu W.P. Adams 77,
123

subsect. 2. Pseudobrathydium R. Keller 78, 80,
112

subsect. Pseudobrathydium sensu W.P.
Adams 77

subsect. 3. Suturosperma R. Keller 78, 80, 81,

113, 123

sect. 14. Oligostema (Boiss.) Stefanoff 76
sect. 12. Origanifolia Stefanoff 76
sect. Pulogensia N. Robson (=sect. 26) 153, 162
sect. 18. Taeniocarpium Jaub. & Spach 76
sect. 23. Triadenioides Jaub. & Spach 76, 83, 84,

141
sect. 30. Trigynobrathys (Y. Kimura) N. Robson 77,

88

sect. 21. Webbia (Spach) R. Keller 82, 90, 133
abilianum N. Robson (Adm2) 90, 174*, 175, 177
aciferum (Greuter) N. Robson (Ads3) 85, 153
acutum Moench 90
adpressum W.P.C. Barton (M21) 119, 120

var. fastigiatum (Elliott) Torrey & Gray
(=M21) 120

var. spongiosum B.L. Rob. (=M21) 120

forma spongiosum (B.L.Rob.) Fernald

(=M21) 120

aegusanum Tineo ex Lojac. (=Adm7) 182
aegypticum L. (Adsl) 84, 85, 147, 148, 149, 152,

153

sensu Braun-Blanquet & Maire (=Adsla) 150

sensu auct. (=Adslb) 150

subsp. aegypticum (Adslc) 150, 152

subsp. maroccanum (Pau) N. Robson
(Ads la) 84, 150, 151*

subsp. webbii (Spach) N. Robson (Adslb) 150

var. maroccanum Pau (=Adsla) 150
aethiopicum Thunb. (Adm6) 90, 169, 170, 177,

179, 180

sensu Jacot Guill. pro parte (=Hu9) 168

sensu Sond. pro parte (=Adm6a) 180

subsp. aethiopicum (Adm6b) 180, 181

subsp. sonderi (Bredell) N. Robson (Adm6a) 90,
180

var. glaucescens Sond. (=Adm6a) 180

var. huillense Engl. (Adm6a) 180
afromontanum Bullock (=Adml8c) 202
afrum Lam. (Adm5) 76, 90, 174*, 178, 179
ambiguum Elliott (=M6) 102
amoenum Pursh (=M1) 94
annulatum Moris (AdmlS) 90, 91, 198, 199

sensu Cufod. (=Adml8a) 199

sensu Milne-Redhead (=Adml8c) 202

STUDIES IN HYPERICVM

215

subsp. afromontanum (Bullock) N. Robson
(AdmlSc) 90, 199, 200*, 201, 202, 203

subsp. annulatum (Adml8b) 91, 199, 201, 202,
203, 204, 205

subsp. degenii (Bornm.) Hayek (=Adml8b) 201

subsp. intermedium (Steud. ex A. Rich.) N.
Robson (AdmlSa) 89, 90, 91, 199, 201
apocynifolium Small (Ml 6) 77, 78, 81, 113, 1 14,

116

arborescens Chapm. (=M14) 112
x arnoldianum Rehd. (=M5x) 101, 102
aspalathoides Willd. (=M13) 1 1 1

sensu Elliott (=M 11) 109

sensu Jennings (=M9a) 108

sensu Small pro parte (=M7) 104, 105
athoum Boiss. & Orph. (Adm20) 91, 203, 204
atlanticum Coss. (=Adml4) 192
atomarium Boiss. (Adm21) 91, 204, 205, 206

sensu Osorio-Tafall & Seraphim (=Adm23) 207

sensu Velen. (=Adml8b) 201
attenuatum sensu Hayata (=Hu6(i)) 162
aureum W. Bartram (=M 1 ) 94
aviculariifolium Jaub. & Spach 76
axillare Lam. (=M6) 102, 104
balfourii N. Robson 84, 147
beccarii N. Robson (Hu7) 87, 162, 164

subsp. beccarii (Hu7a) 86, 165

subsp. steenisii N. Robson (Hu7b) 86, 87, 165
bifurcatum N. Robson (Hu4) 86, 159, 168
bissellii B.L. Rob. (=M24) 123, 124
bojerianum sensu H. Perrier pro parte (=Hu9) 168
bonaparteae W.P.C. Barton (=M21) 120
brachyphyllum (Spach) Steud. (Mil) 80, 105, 106,

107*, 109

Brathydium group (M23-24) 78
brathydium Steud. (=M22) 121
buckleyi M.A. Curtis (Ml 5) 77, 78, 80, 112, 113
canadense van oviforme R. Keller (=M22?) 121
canariense L. (Wl) 82, 90, 134, 135, 136*, 137, 140

[van] fifloribundum (Alton) Bornm. (=W1) 135

var. montanum Buch (=W 1 ) 1 34

[van] y platysepalum (Spach) Bornm.
(=W1) 135

[var.] 7 salicifolium Choisy (=W1) 134

[var.] P triphyllum Choisy (=W1) 134

[var.] a typicum Bornm. (=W 1 ) 135
canescens Trevir. (?=Adm9) 186
caprifolium group (Adml3-15) 89, 21 1
caprifolium Boiss. (AdmlS) 89, 90, 191*, 192, 193

subsp. naudinianum (Coss. & Durieu) Maire

(=Adml4) 192

carbonelli Sennen & Mauricio (=Adm9) 186, 187
cardiophyllum Boiss. (Ar3) 138, 139*, 140, 141
chamaenerium (Spach) Steud. (=M20) 118
chapmanii W.P. Adams (Ml 4) 112
ciliatum var. pseudociliatum R. Keller

(=Adml7) 197
cistifolium Lam. (M18) 77, 78, 82, 1 14, 115*, 116,

117, 118, 119

sensu Coulter pro parte (=M 17) 114

sensu Coulter pro parte (=M20) 1 1 8
coadunatum Chr. Sm. (Adml3) 89, 90, 189, 190,

191*, 192,211

sensu Maire (=Adml4) 192

var. atlanticum Ball (=Adml4) 192

var. disjunctum R. Keller (=Adml3) 189
collenettiae N. Robson (Adml 1) 90, 183*, 188, 189
confertum Moench (=Adml7) 196
conjunction N. Robson (=Adm3) 177

sensu Agnew (=Adm4) 178
conjungens N. Robson (Adm3) 90, 174*, 177, 178,

179, 180

const anzae Urban 133
coris? sensu Walter (=M7) 104
corymbosum Moench (=W1) 134
creticum Hort. ex Link (=Adsl) 149
crux-andreae (L.) Crantz (M25) 78, 82, 124, 126*.

127, 128, 129, 130, 131

cryptopetalum Vogel (=M2) 96

cubense Turcz. (=M9a) 106

cuisinii Barbey (Adm22) 89, 91, 200*, 205, 206

cuneatum Poir. (=T5) 146

[var.] b. fragile Post (=T5) 146

var. maximum Post (=T5) 146

[van] c. pallidum Post (=T5) 146
x dawsonianum Rehd. (=M4x) 100
debile Sa\\sb. (=W1) 134
decaisneanum Coss. & Daveau (Adm24) 90, 208
degenii Bornm. (=Adml8b) 201
delphicum Boiss. & Heldr. (Adml 9) 91, 202, 203,

204

densiflorum group (M 4—8) 98
densiflorum Pursh (M5) 79, 80, 81, 97, 98, 100,

101, 104

var. lobocarpum (Gatt.) Svenson (=M4) 99
densiflorum x galioides? (M5x) 101
densiflorum x katmianum sensu Rehd. (=M5) 102
densiflorum x kalmianum? (=M5xx?) 102
densiflorum x lobocarpum (=M5x?) 102
densiflorum x prolificum (5xx) 101, 102
diosmoides Griseb. 133
dolabriforme Vent. (M24) 78, 82, 122, 123, 124
edisonianum (Small) W.P. Adams & N. Robson

(M27) 82, 126*, 128, 129
elegans Stephan ex Willd. 179
elegantissimum Cr. (=Adml7) 196
ellipticum Hook. (M22) 79, 82, 120, 122

forma foliosum Viet. (=M22) 121, 122

forma submersum Fassett (=M22) 121, 122
elodeoides Choisy 76
elodes L. (El) 76, 88, 89, 90, 191*, 209, 21 1

forma glabratum Druce (=E1) 209
empetrifolium sensu Kotschy ex Jaub. & Spach

(=Ads2) 152

exile W.P. Adams (=M9c) 108
fasciculatum group (Ml 3- 14) 78, 81, 98, 106
fasciculatum Lam. (M13) 81, 107*, 110, 112
fasciculatum Michx. ex Willd. (=M6) 102

sensu Alain, pro parte (=M9a) 108

sensu Alain, pro parte (=M9c) 108

sensu Choisy, pro parte (=M7) 104

sensu Coulter, pro parte (=M1 1) 109

sensu Torrey & Gray, pro parte (=M9) 106

var. aspalathoides (Willd.) Torrey & Gray
(=M13) 111

sensu Chapm. (=M11) 109

sensu Torrey & Gray, pro parte (=M7) 104

var. laxifolium Choisy, pro parte (=M7) 104

[var.] P sensu Choisy, pro parte (=M6) 102

[var.] P sensu Choisy, pro parte (=M7) 104
fastigiatum Elliott (=M21) 120
fieriense N. Robson (Tl) 84, 142, 143, 144*
floribundum Alton (=W\) 134, 135, 137
foliosum sensu Brouss. ex Webb & Berth.

(=Adml6) 193

sensu Jacq. (=M2) 96, 101
formosissimum Takht. 76
frondosum Michx. (Ml) 78, 79, 80, 82, 94, 95*,

98, 101, 118, 123, 124, 127
frondosum x prolificum (Mix) 96
fulgidumRaf. (=M13?) Ill
galioides group (M6-8) 78
galioides Lam. (M6) 78, 81, 101, 102, 103*. 104,

106

sensu Griseb., pro parte (=M9c) 108

sensu Sauvalle (=M9c) 108

var. ambiguum (Elliott) Chapm. (=M6) 102, 104

var. axillare (Lam.) Griseb. (=M6) 102

sensu Griseb. pro parte (=M9c) 108

var. cubense Griseb. (=M9c) 108

var. fasciculatum (Lam.) Svenson (=M13) 1 1 1

var. fasciculatum (Lam.) Svenson, pro parte
(=M9) 106

var. lloydii Svenson (=M8) 105

[var.] pallidum Mohr (=M6) 102

var. reductum Svenson (=M7) 104

galioides x lobocarpum sensu Rehd. (=M5x?) 101
glandulosum Alton (Adml) 89, 90, 172, 174*, 175,

184, 194

glandulosum Gilib. (=Adml7) 196
glaucum Michx. (=M23) 122
glomeratum Small (=M5) 100
gracile Boiss. (=Adm23) 207
habbemen.se A.C. Sm. (=Hu5) 160, 162
hayataeY. Kimura (=Hu6(i)) 162, 163
hellwigii Lauterb. (=Hu5) 159
helodes St.-Lager (=E1) 209

forma terrestre Cluck (=E1) 209, 21 1
helodeum St.-Lager (=E1) 209
hengshanense W. T Wang 76
heterophyllum Vent. (Hel) 147, 151*
hetemstylum Part. (=Adsl) 149
hirsutum Asso (=Adml5) 192
hirsutum L. 198

sensu Sibth. & Sm. (=Adm21) 204
huber-morathii N. Robson 76
humbertii sensu Spirlet, pro parte (=HulO) 169
hypericoides (L.) Crantz (M29) 78, 82, 124, 129,

130

subsp. hypericoides (M29a) 82, 130, 131, 132,
133

subsp. multicaule (Michx. ex Willd.) N. Robson
(M29b) 82, 131, 132

subsp. prostratum N. Robson (M29c) 78, 82,
131, 133

var. hypericoides (=M29a) 131

var. multicaule (Michx. ex Willd.) Fosberg
(=M29b) 131, 132

var. multicaule (Michx. ex Willd.) Waterfall

(=M29b) 132
imbricatum Poulter 76
interior Small (=M5) 81, 100, 101
intermedium Steud. ex A. Rich. (=Adml8a) 199

forma obtusifolium R. Keller (=Adml8a) 201
isophyllum Steud. (=M19) 1 17
japonicum sensu Warb. (+Hu5) 159

var. pinnatinervium Bakh. f. (=Hu7) 164
x joerstadii Lid (Admix) 89, 90, 175, 194
kalmianum L. (M3) 80, 95*. 98, 101

var. majus Gatt. (=M2) 97
kalmianum x densiflorum'? (=M5) 99
kalmianum x prolificum (M3x) 99
kiboense Oliv. (Adm4) 90, 174*. 177, 178, 179

sensu N. Robson, pro parte (=Adm2) 175
kiloense sensu H.H. Johnston (=Adm4) 178
kunaianum Gilli (=Hu5) 160
lanceolatum Lam. 85

subsp. angustifolium (Lam.) N. Robson 86, 157
lanuginosum Lam. (Adm23) 89, 91, 200, 205. 206,

208

sensu d'Urv. (=Adm21) 204

subsp. atomarium (Boiss.) Holub
(=Adm21) 205

subsp. gracile (Boiss.) Holmboe ex J. Thiebaut
(=Adm23) 207

subsp. millepunctatum Holmboe (=Adm23) 207

[var.] P gracile (Boiss.) Boiss. (=Adm23) 207

var. pestalozzae (Boiss.) N. Robson
(=Adm23) 92, 207, 208

var. scabrellum (Boiss.) N. Robson

(=Adm23) 92, 207, 208
ligustrinum Pursh (=M 17) 114
limosum Griseb. (Ml 0) 106, 108, 109
lissophloeus W.P. Adams (M12) 78, 107*, 110
lloydii (Svenson) W.P. Adams (M7) 80. 81, 103*.

105, 106

lobocarpum Gatt. (M4) 80, 81, 98, 99, 100, 101
lobocarpum x prolificum (M4x) 100
lusitanicum Poir. (=Adm9) 185
macgregorii F. Mull. (Hu3) 86, 158, 159

sensu Hoogl. (=Hu2) 157

sensu Lauterb. (=Hu5) 159

subsp. punctatum N. Robson (=Hu2) 157
maculatum Cr. 90

216

N.K.B. ROBSON

madagaseariense (Spach) Steud. 85

maritimum Sieber (=Adslb) 150, 152

michauxii Poir. (=M6) 102

microsepalum (Torrey & Gray) A. Gray ex S.
Watson (Ml 9) 77,78,82, 113, 115*, 117, 118

milne-redheadii Gilli (=Adm3) 177

minutum Poulter 76

montanum group (spp. 17-24)

montanum L. (Adml7) 89, 90, 194, 195M98, 199
subsp.? elegantissimum (Cr.) G. Jav.

(Adml7) 197

Ivar.l P caucaticum Boiss. (=Adml7) 196, 198
var. maculantherum Sagorsky 198
var. pilosum Horwood 198
var. punctatum Andreansky 198
var. scaberulum G. Beck (=Adml7) 197, 198
[var.l P scabrum Koch (=Adml7) 196. 198
| van] P triphyllum Choisy (=Adml7) 196
[var.l « typicum G. Beck (=Adml7) 197, 198
[var.J P sensu Ledeb. (=Adml7) 196
forma abbreviatum Reinecke (=Adml7) 197
forma humifusoides Kuntze (=Adml7) 197
forma tubpmlificum Murr (==Adm) 197
forma ternatum Borbas (=Adml7) 197

multicaule Lam. 132

myrtifolium Lam. (M23) 77, 78, 82, 122, 123, 124

mvrtilloidet Fenzl (=T5) 146

nagasawai Hayata (Hu 6(i)) 86, 154, 162, 163, 164
var. nit-rum Y. Kimura (=6(i)) 162
var. typicum Y. Kimura (=6(i» 162

nanum Poir. (Ar4) 139*. 140, 141
var. nanum (Ar4a) 140
var. prostratum Boiss. (Ar4b) 141
var. uniflorum Bornm. (=Ar5) 141

natalense J.M. Wood & M.S. Evans (Hu8) 86, 87,
165, 167*, 168, 170
var. petiolatum Bredell (=Hu8) 165

naudinianum Coss. & Durieu (Adml4) 89, 90, 190,
191*. 192, 193

nigropunctatum Norlindh (=Hu9) 168

nitidum group or complex (M9-1 1) 78, 81, 98

nitidum Lam. (M9) 78, 81, 104, 106, 112

subsp. cubense (Turcz.) N. Robson (M9a) 106,

107*, 109
subsp. exile (W.P. Adams) N. Robson

(M9c) 106, 108, 109
subsp. nitidum (M9b) 106, 108, 1 12

nokoenseOhwi (Hu6(ii)) 86, 154,163,164

nothum Rehd. (=M5) 100, 102

nudiflorum Michx. ex Willd. (M17) 77, 78, 81,
114, 115*. 116, 117, 118
sensu Rchb. (=M20) 118
sensu auct. pro parte (=M16) 1 13
[var.] P ovatum Choisy (=M17) 1 14
[van] "Y ramosum Choisy (=M17?) 1 14
[var.] P sensu Torrey & Gray (=M16) 1 13

oklahnmense E.J. Palmer (=M4) 99

opacum Torrey & Gray (=M18) 116. 117

origanifolium Willd. 76

pallens Banks & Solander (T5) 84, 144*, 145, 146

palustre Salisb. (=E1) 209

pamphylicum N. Robson & P. Davis (Ar2) 138,
139

papuanum Ridl. (Hu5) 86, 156*. 159, 161, 162, 164

pauciflorum Kunth 129

peltatum sensu Eisner et al. (=M25) 125

peplidifolium A. Rich. (HulO) 86, 167*, 168, 169,
170

var. anagallidifolium Chiov. (=HulO) 169
var. diestelianum Engl. (=HulO) 169
var. oblongifolium Engl. (=HulO) 169
var. ovatum (Engl.) Engl. (=HulO) 169
forma humile Riva (=Hu 10) 169
var. mbuttum Baker f. (=HulO) 169
forma ovatum Engl. (=HulO) 169
forma parvifolium Engl. (=HulO) 169
forma mbustum (Baker f.) Engl. (=HulO) 169

perfoliatum L. 198

var. annulalum (Moris) Fiori (=Adml8b) 201

var. pseudociliatum (R. Keller) Woron.

(=Adml7) 197

perfoliatum Munby (=Adml4) 190
perforatum L. 90

sensu Poir. (=Adm5) 178
pestalozzae Boiss. (=Adm23) 206
platypetalum (Spach) Steud. (=W1) 135
platysepalum (Spach) Walp. (=W1) 135
procumbent Desf. ex Willd. (=M24) 123
procumbens Michx. (=M24) 123
prolificum L. (M2) 78, 80, 81, 95*, 96, 97, 98, 101,

106, 113, 114

sensu Torrey & Gray, pro parte (=M5) 100

var. aureum (W. Bartram) Koehne (=M1) 94

var. densiflorum (Pursh) A. Gray (=M5) 100

var. montanum Gatt. (=M2) 97
psilophytum (Diels) Maire (Adm8) 90, 184, 185
pubescens Boiss. (Adm7) 90, 181, 183*, 184, 185,

197, 188, 190

pubescens x tomentosum (Adm7x) 184
pulogense Merr. (Hu6) 86, 87, 154, 162, 163, 164
pumillum Sesse & Mocino 129
punctulosum Bertol. (=M18) 116
quartinianum A. Rich. 79, 90, 175
randaiense Hayata (=Hu6(i)) 162, 163
reductum (Svenson) W.P. Adams (=M7) 104, 105
reflexum L. f. (Adml6) 89, 90, 175, 193, 194,

195*. 198

var. lanuginosum Pitard (=Adml6) 194

var. leiocladum Bornm. (=Adml6) 193, 194

var. myrtillifolium Bornm. (=Adml6) 175, 193,

194
revolutum Vahl 135, 146

subsp. revolutum 90
revolutum R. Keller (=M5) 100
roeperianum W.G. Schimper ex A. Rich. 82, 135,

175
rosmarinifolium Lam. (=M18) 116

sensu Choisy (=M23) 122

sensu Torrey & Gray (=M5) 100
rosmarinifoliuml sensu Elliott (=M6) 102
mstratum Raf. (=M4?) 99
rugelianwn Kunze (=M1) 94
rupestre Jaub. & Spach (Arl) 83, 137, 138, 139*,

140, 141

sensu H. Perrier, pro parte (=Hu9) 168

[van) a rotundifolium Jaub. & Spach (=Arl ) 137

[var.l P ovalifolium Jaub. & Spach (=Arl) 137
russeggeri (Fenzl) R. Keller (Ads2) 85, 147, 152,

153

salsugineum N. Robson & Hub.-Mor. 76
sanctum Degen (=Adm20) 203
saruwagedicum Diels (Hu2) 156*, 157, 158, 159
scabrellum Boiss. (=Adm23) 207
scopulorum Balf. f. (T2) 83, 84, 142, 143, 144*,

145

sessiliflorum Willd. ex Spreng. (=M23) 122
sewense N. Robson (Hul) 86, 87, 154, 156*, 157
sieberi (Spach) Nyman (=Adslb) 150
silenoides Juss. subsp. minus N. Robson 76
sinaicum Hochst. ex Boiss. (Adml2) 90, 183*.

188, 189
socotranum Good 83, 84, 145, 147

subsp. socotranum 83

somaliense N. Robson (AdmlO) 90, 187, 188
tonderi Bredell (=Adm6a) 180

var. transvaalense Bredell (=Adm6a) 180
sp. A sensu Milne-Redh. (=Adm4) 178
sp. B sensu Milne-Redh. (=Adm3) 177
sp. C sensu Milne-Redh. (=HulO) 169
sp. aff. sinaicum sensu Collen. (=Adml 1) 188
spachianum Steud. (=Adslb) 150
spathulatum R. Keller (=M6) 102
spathulatum (Spach) Steud. (=M2) 97, 132
sphaerocarpum Michx. (M20) 78, 82, 1 17, 118,

119, 120, 122, 124

sensu W.P.C.Barton (=M22) 121

var. sphaerocarpum Svenson (=M20) 1 18
var. turgidum (Small) Svenson (=M20) 118, 119
splendent Small (=M1) 94
spruneri Boiss. 198

slant (Michx.) Adams & Robson (=M25) 125
stragulum Adams & Robson (=M29b) 131, 132
suberoswn Salzm. ex Boiss. (=Adm9) 185

var. psilophytum Diels (=Adm8) 184
suffruticosum W.P. Adams & N. Robson (M28) 78,

82, 126*, 127, 128, 129
supinum Vis. (=Adm9) 186
supinum Vis., pro parte (=Adm 21) 204
supinum tomentosum majus & hispanicum Bauhin

(=Adm9) 185
supinum tomentosum minus & monspeliacum

Bauhin (=Adm9) 185
Suturosperma group (M 16-22) 78
suzukianumY. Kimura (=Hu6(i) 162, 163
synstylum N. Robson 78, 79, 90
taiwanianium Y. Kimura (=Hu6(i» 162, 163
var. ohwii\. Kimura (=Hu6(i)) 162
var. taiwanianum (=Hu6(i)) 162
taubertii Asch. & Barbey ex Coss. (=Adm24) 208
tauricum sensu hort. ex Ledeb. (=Adml7) 196
tenellum Kotschy ex Boiss. (=T5) 146
tenuifolium Pursh (M8) 80, 81, 103*, 104, 105
ternatum Poulter (T4) 84, 144*, 145, 146
tetrapetalum Lam. (M26) 78, 82, 126, 127, 128

[var.] P sensu Lam. (=M25) 124
tetrapterum Fr. 90

tomentosum L. (Adm9) 90, 183*, 184, 185, 187
sensu Decne. (=Adml2) 189
sensu Durand(=El) 209
sensu Guss. (=Adm7) 182
subsp. carbonelli Sennen & Mauricio

(=Adm9) 186

subsp. eu-tomentosum Maire (=Adm9) 186
var. carbonelli (Sennen & Mauricio) Maire
(=Adm9) 186

subsp. lusitanicum (Poir.) Willk. (=Adm9) 186
subsp. psilophytum (Diels) Maire (=Adm8) 184
subsp. pubescens (Boiss.) Ball (=Adm7) 181
var. damnatorum Maire (=Adm7) 182, 184
var. viridulum Pau (=Adm7) 182, 184
subsp. wallianum Maire (=Adm9) 186
[van] P ambiguum Perez Lara (=Adm7x) 184
var. densiflorum Sennen (=Adm) 186
var. dissitiflorum Roemer (=Adm9) 186
| var. | a genuinum Perez Lara (=Adm9) 186

subvar. elevatum Perez Lara (?=Adm7x) 184
var. glabrescens Porta (=Adm9) 186
var. intermedium Coss. ex Willk. (=Adm7x) 90,

184
[var.] "Y lusitanicum (Poir.) Perez Lara

(=Adm9) 186

var. palustre Batt. (=Adm9) 186
[var.] y pubescens (Boiss.) Perez Lara

(=Adm7) 181

var. racemosum Batt. (=Adm9) 186
var. ramosissimum Sennen (=Adm9) 186
tomentosum x pubescens (Adm7x) 90
tortuosum Balf. f. (T3) 84, 142, 143, 144*, 145
triplinerve Vent. (=M6) 102
turgidum Small (=M20) 1 18
undulatum Schousb. ex Willd. 179
vacciniifolium Hayek & Siehe (Ar5) 138, 139*,

141

x vanjleetii Rehd. (=Mlx) 96
vesiculosum Griseb. 198
webbii (Spach) Steud. (=Adslb) 150
wilmsii R. Keller (Hu9) 86, 87, 167*, 168, 169,

170

woodii R. Keller (=Hu8) 165
yezoense Maxim. 154
Isophyllum Spach (= sect. 20 subsect. 3) 77, 113

drummondii Spach (=M 19) 77, 113, 117
Myriandra adpressa (W.P.C. Barton) K. Koch
(=M21) 120

STUDIES IN HYPERICUM

217

hrachyphylla Spach (=MI 1) 109

brathydis Spach (=M13) 111

galioides (Lam.) Spach (=M6) 102

xliiuca (Michx.) Spach (=M23) 122

ledifolia Spach (=M2) 96

mii-hauxii (Pair.) Spach (=M6) 102

nitida (Lam.) Spach (=M9) 106

nudiflora (Michx. ex Willd.) Spach (=M17) 1 14

pmlifica (L.) Spach (=M2) 96
var. ramosa K. Koch (=M6) 102
var. spathulata (Spach) K. Koch (=M2) 97

spathulata Spach (=M2) 96
Nor\xca kalmiana (L.) K. Koch (=M3) 98

Pinus nigra J.F. Arnold subsp. pallasiana (Lamb.)

Holmboe 147

Spachelodes eludes (L.) Y. Kimura (=E1) 209
Streptalon dolabriforme (Vent.) Rat'. (=24) 123
Triadenia aegyptica (L.) Boiss. (=AdsI ) 149

var. micmphylla (Spach) Maire (=Adsl) 149
maritima (Sieber) Boiss. (=Adslb) 150
subsp. weissii Stamatiadou (=Adslb) 150
[var] B. webbii (Spach) Hayek (=Adslb) 150
micmphylla Spach (=Adsl) 149
russeggeri Fen/.l (=Ads2) 152
sieberi ; Spach (=Adslb) 150
thymifolia Spach (=Adslb) 150

webbii : Spach (=Adslb) 150
Tripentas helodex (L.) Asch. & Graebn. (=E1) 209
Webhia canariensis (L.) Webb & Berth. (=WI ) 135
[var.j fyfloribunda (Aiton) Pitard &

Proust (=W1) 135
[var.] Y platypetala (Spach) Pitard & Proust

(=W1) 135
[var.] a typica (Bornm.) Pitard & Proust

(=WI) 135

fluribunda (Aiton) Spach (=WI) 134, 135
hetemphylla Spach (=W I) 134, 135
platypetalu Spach (=W1) 135
platysepala Spach (=W 1 ) 1 35

Bulletin of The Natural History Museum
Botany Series

Earlier Botany Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is not shown,
this may also be obtained from the same address.

Volume 19

A new species of Maytenus (Celastraceae) in Ethiopia. Sebsebe

Demissew. 1989. Pp. 1-3, 1 fig.

Central American Araliaceae - a precursory study for the Flora

Mesoamericana. M.J. Cannon & J.F.M.Cannon. 1989. Pp. 5-

6 1,36 figs.

A revision of the Solarium nitidum group (section Holophylla

pro parte): Solanaceae. S. Knapp. 1989. Pp. 63-102, 21 figs.

Six new species of Solatium sect. Geminata from South

America. S. Knapp. 1989. Pp. 103-1 12, 8 figs.

The application of names of some Indian species of Ocimum

and Geniosporum (Labiatae). J.R. Press & V.V. Sivarajan.

1989. Pp. 11 3- 11 6, 4 figs.

Revision of Piper (Piperaceae) in the New World 1 . Review of

characters and taxonomy of Piper section Macrostachvs. M.C.

Tebbs. 1989. Pp. 1 17-158, 41 figs. £48.00

Volume 20

No. 1 Studies in the genus Hypericum L. (Guttiferae) 8. Sections 29.

Brathys (part 2) and 30. Trigynobrathvs. N.K.B. Robson. 1990.
Pp. 1 - 1 5 1 , 22 figs, 46 maps. £45.00

No. 2 The marine algal flora of Namibia: its distributions and

affinities. G.W. Lawson, R.H. Simons and W.E. Isaac. 1990.

Pp. 153-168, 1 fig, 7 plates.

The infrageneric classification of Gentiana (Gentianaceae). T-

N. Ho and S.-W. Liu. 1990. Pp. 169-192, 13 figs.

Revision of Piper (Piperaceae) in the New World. 2. The

taxonomy of Piper section Churumavu. M.C. Tebbs. 1990. Pp.

193-236, 49 figs. £25.00

Volume 21

No. 1 Historical and taxonomic studies in the genus Titanoderma

(Rhodophyta, Corallinales) in the British Isles. Y.M. Chamber-
lain. 1991. Pp. 1-80, 247 figs.

No. 2 Early collections of the Holy Thorn (Crataegus monogyna cv.
Biflora). A.R. Vickery. 1991. Pp. 81-83, 1 fig.
A taxonomic study of the species referred to the ascomycete
genus Leptorhaphis. B. Aguirre-Hudson. 1991. Pp. 85-192, 76
figs.

The typification and identification of Calymperes
crassilimbatum Renauld & Cardot (Musci: Calymperaceae).
L.T. Ellis. 1991. Pp. 193-194, 1 fig.

Volume 22

No. 1 An account of southern Australian species of Lithophyllum

(Corallinaceae, Rhodophyta). Wm. J. Woelkerling and S.J.
Campbell. 1992. Pp. 1-107, 63 figs. £37.00

No. 2 Palynological evidence for the generic delimitation of Sechium
(Cucurbitaceae) and its allies. J.L. Alvarado, R. Lira-Saade &
J. Caballero. 1992. Pp. 109-121.

Seaweeds of the western coast of tropical Africa and adjacent
islands: a critical assessment. IV. Rhodophyta (Florideae) 3.
Genera H-K. J.H. Price, D.M. John & G.W. Lawson. 1992. pp.
123-146.

Two new species of Solatium section Geminata (Solanaceae)
from Cerro del Torr in western Colombia. S. Knapp. 1992. Pp.
147-152.

Fissidens ceylonensis Dozy & Molkenb. (Musci:
Fissidentaceae) and some allied taxa from southern India. L.T.
Ellis. 1992. Pp. 153-156, 2 figs.

New species of Piper (Piperaceae) from Central America. M.
Tebbs. 1992. Pp. 157-158.

Studies on the Cretan flora 1 . Floristic notes. N.J. Turland.
1992. Pp. 159-164.

Studies on the Cretan flora 2. The Dianthus juniperinus
complex (Caryophyllaceae). N.J. Turland. 1992. Pp. 165-169.
£37.50

Volume 23

No. 1 Revision of Piper (Piperaceae) in the New World 3. The

taxonomy of Piper sections Lepianthes and Radula. M.C.
Tebbs. 1993. Pp. 1-50, 18 figs.

Mounting techniques for the preservation and analysis of
diatoms. S.J. Russell. 1993. Pp. 51-54. 1 fig. £37.50

No. 2 New taxa of Gentiana (Gentianaceae) from Western China and
the Himalayan region. T.-N. Ho and S.-W. Liu. 1993. Pp. 55-
60. 2 figs.

New combinations, names and taxonomic notes on Gentianella
(Gentianaceae) from South America and New Zealand. T.-N.
Ho and S.-W. Liu. 1993. Pp. 61-66.
Studies in Hypericum: validation of new names. N.K.B.
Robson. 1993. Pp. 67-70.

Generic monograph of the Asteraceae-Anthemideae. K.
Bremer and C.J. Humphries. 1993. Pp. 71-177. 12 figs. £37.50

Volume 24

No. 1 Pre-Linnaean references for the Macaronesian flora found in

Leonard Plukenet's works and collections. J. Francisco-Ortega,

A. Santos-Guerra and C.E. Jarvis. Pp. 1-34.

Studies on the lichen genus Sticta (Schreber) Ach.: II.

Typification of taxa from Swartz's Prodromus of 1788. D.J.

Galloway. Pp. 35^8.

Seaweeds of the western coast of tropical Africa and adjacent

islands: a critical assessment. IV. Rhodophyta (Florideae) 4.

Genera L-O. D.M. John, G.W. Lawson, J.H. Price, W.F.

Prud'homme van Reine and W.J. Woelkerling. Pp. 49-90.

Studies on the Cretan flora 3. Additions to the flora of

Karpathos. N.J. Turland and L. Chilton. Pp. 91-100.

No. 2 Observations on the benthic marine algal flora of South

Georgia: a floristic and ecological analysis. D.M. John, P.J.A.

Pugh and I. Tittley. Pp. 101-1 14.

Studies in Pseudocyphellaria (Lichens) IV. Palaeotropical

species (excluding Australia). D.J. Galloway. Pp. 1 15-160.

Morphology and ecology of seedlings, fruits and seeds of

Panama: Bixaceae and Cochlospermaceae. N.C. Garwood. Pp.

161-172.

A study of Bixa (Bixaceae), with particular reference to the leaf

undersurface indumentum as a diagnostic character. R.E.

Dempsey and N.C. Garwood. Pp. 173-180.

Volume 25

No. 1 A revision of Rutilaria Greville (Bacillariophyta). R. Ross. Pp.

1-94.
William Roxburgh's St Helena plants. Q.C.B. Cronk. Pp. 95-98.

No. 2 Seaweeds of the western coast of tropical Africa and adjacent
islands: a critical assessment. IV. Rhodophyta (Florideae) 5.
Genera P. G.W. Lawson, W.J. Woelkerling, J.H. Price, W.F.
Prud'homme Van Reine and D.M. John. Pp. 99-122.
A new species of Odontorrhynchos (Orchidaceae,
Spiranthinae) from Boliva.D.L. Szlachetko. Pp. 123-125.
Linnaeus's interpretation of Prospero Alpino's De plantis
exoticis, with special emphasis on the flora of Crete. N.J.
Turland. Pp. 127-159.
Book review. M.G. Gilbert. P. 161.

Volume 26

No. 1 A morphological study of Chaetoceros species

(Bacillariophyta) from the plankton of the Pacific ocean of
Mexico. D.U. Hernandez-Becerril. 1996. Pp. 1-73.

CONTENTS

75 Studies in the genus Hypericum L. (Guttiferae) 6. Sections 20. Myriandra to 28. Elodes
N.K.B. Robson

jlletin of The Natural History Museur
BOTANY SERIES

Vol. 26, No. 2, November 1996