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Bulletin of the 

British Museum (Natural History) 



BRITISH MUSEUM 
(NATURAL HISTORY) 

25 FEB 1988 



PRESENTED 
GENERAL LIBRARY 



Botany series Vol 16 1987 



British Museum (Natural History) 
London 1987 



Dates of publication of the parts 



Nol 

No 2 
No 3 
No 4 



ISSN 0068-2292 



\ 



29 January 1987 

25 June 1987 
27 August 1987 

26 November 1987 



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



Contents 

Botany Volume 16 

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

No 2 The Hchen genus Ramalina in Australia 

G.N.Stevens 107 

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

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



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-3FEBI987 



Bulletin of the i 

British Museum (Natural History) 



Studies in the genus Hypericum L. 

(Guttiferae) 

7. Section 29. Brathys (part 1) 

Norman K. B. Robson 



Botany series Vol 16 No 1 29 January 1987 



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

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WorWLwf abbreviation: Bull. Br. Mus. not. Hist. (Bot.) 



© Trustees of the British Museum (Natural History), 1987 



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

Editor of Bulletin: Mr J. R. Laundon 

Assistant Editor: Dr A. J. Harrington 

Editor's Assistant: Miss M. J. Short 



ISBN 565 08014 8 
ISSN 0068-2292 

British Museum (Natural Hist 
Cromwell Road 
London SW7 5BD 




/| Botany series 

^ Vol 16 No 1 pp 1-106 



Issued 29 January 1987 



(NATURAL KIS^ORV) 

»3FL '757 



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

Norman K. B. Robson 

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

Contents 

Synopsis 1 

1 . Introduction 2 

Morphology 3 

(a) Leaves 3 

(b) Inflorescences 5 

(c) Flowers and fruits 5 

Cytology and hybrids 7 

Distribution and evolution 7 

(a) Wide disjunctions 7 

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

(Spp.3-23) 9 

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

secretions (Spp. 24-26,61) 10 

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

secretions (Spp. 27-30) 10 

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

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

(f) Species with pseudopetiolate leaves deciduous above the base 

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

2. Systematic treatment 12 

3 . Acknowledgements 103 

4. References 104 

5. Systematic index 104 

Synopsis 

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

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

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



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



2 NORMAN K. B. ROBSON 

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

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

1. Introduction 

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

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

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

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

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

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

Tridia Korth. in Tijdschr. Naturl. Gesch. Physiol. 3: 17(1836). 

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

Sarothra sect. Spachium sehts Japonica Y. Kimura, tom. cit. : 233 (1951). 
Type: //. mutilum L. (Y. Kimura, 1951, lectotype). 

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

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



THE GENUS HYPERICUM L. 3 

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

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

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

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

Kimura). 

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

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

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

monochasial sect. 29. Brathys 

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

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

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

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

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

decurrent or sometimes perfoliate sect. 31 . Trigynobrathys 

Morphology (Fig. 2) 

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

(a) Leaves 

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

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



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THE GENUS HYPERICUM L. 5 

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

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

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

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

(b) Inflorescences 

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

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

(c) Flowers and fruits 

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

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

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



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THE GENUS HYPERICUM L. 7 

Cytology and hybrids 

Very little cytological work has been done on the major part of Sect. Brathys, the only recorded 
chromosome count for a woody species being n = 12 for 7 //. irazuense (Part 2: table 7) . As this is 
the primitive number for the whole genus (Part 2: 150), there is no available information about 
possible chromosome variation within the woody members of the section. The numbers n = 6, 
12, 24 have been recorded for herbaceous species (see part 8). 

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

Distribution and evolution (Fig. 3) 

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

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

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

(a) Wide disjunctions 

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



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

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

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

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

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

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

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



10 NORMAN K. B. ROBSON 

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

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

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

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

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

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

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



THE GENUS HYPERICUM L. 11 

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

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

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

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

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

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

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

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



12 NORMAN K. B. ROBSON 

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

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

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

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

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

2. Systematic treatment 

Sect. 29. BRATHYS (Mutis ex L.F.) Choisy 
Prodr. Monogr. Fam. Hyperic: 3S, 58 (1821).* 

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

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



THE GENUS HYPERICUM L. 13 

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

Basic chromosome number (x): 12; ploidy 1,2,4. 

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

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

80 species (+8 subspecies). 

Key to sect. 29. Brathys 

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

interfoliar ridges 2 

Leaves persistent (falling with cortex) or deciduous above base; stem articulate and/or with 
interfoliar ridges 5 

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

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

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

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

outcurving 2a. styphelioides subsp. clarense{p. 21) 

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

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

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

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

branches often absent or sparse 6 

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

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

internodes and leaves falling as one 7 

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



14 NORMAN K. B. ROBSON 

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

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

diam 54. millefolium (p. 93) 

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

Leaves united by interfoliar ridge or lamina proper 22 

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

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

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

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

internodes smooth 11 

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

corky emergences 13 

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

shrubto3mtall 4. garciae(p. 26) 

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

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

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

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

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

13(10) Leaves with 3 basal veins, subcoriaceous 14 

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

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

Leaves spreading to appressed before falHng 15 

15(14) Leaves deciduous 16 

Leaves persistent 18 

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

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

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

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

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

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

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

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

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

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

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

Leaves with margin incurved, closely appressed, subpersistent 

3c. p/ie//os subsp. platyphyllum(p. 25) 

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

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

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

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

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

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

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

mm long 24 



THE GENUS HYPERICUM L. 15 

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

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

11. papillosum {p. 34) 

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

lycopodioides) 26 

Leaves with bases of lamina united, often becoming deflexed 32 

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

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

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

12. lycopodioides {p. 36) 
Leaves chartaceous, weakly or not tetrastichous, apex acute to rounded, margin usually 
undulate 13. thuyoides(p. 37) 

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

Leaves shortly petiolate or base angustate, if base narrowly cuneate, then lamina linear to 
acicular 30 

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

22. hartwegii(p. 51) 
Leaves 2-4 mm long, narrowly elliptic or narrowly oblong to linear; sepals 0-7-1-7 mm 
wide, apex incurved-cucullate 23. maguirei{p. 52) 

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

19. sabiniforme(p. 45) 
Leaves smooth beneath, base narrowly cuneate to angustate or parallel-sided; sepals acute 
to obtuse 31 

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

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

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

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

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

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

scarcely hyaline 16. cuatrecasii{p. 41) 

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

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

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

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

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

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

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

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

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

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

61. cymobrathys(p. 103) 
Leaves with pinnate venation or 1 -nerved; flowers oo in ± lax cymes; suffrutices or herbs 

[62-80. See Part 8] 

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

61. cymobrathys(p. 103) 
Leaf venation flabellate or pinnate or 1-nerved, veins branched or not; flowers solitary 
(except in 26. mexicanum) 39 



16 NORMAN K. B. ROBSON 

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

linear 40 

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

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

capitate 41 

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

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

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

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

26. mexicanutn(p. 56) 

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

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

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

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

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

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

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

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

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

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

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

46(45) Leaves petiolate or abruptly narrowed at the base ; stem sometimes apparently articulated 47 

Leaves sessile , gradually narrowed or parallel-sided at the base ; stem not articulated 54 

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

petiole 49. jaramilloi{p. 86) 

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

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

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

Stigmas clavate or small ; leaves not glaucous, ± lustrous 50 

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

Leaves with margin or whole lamina incurved 52 

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

51a. caracasanutn suhsp. caracasanum(p. 89) 
Leaves 8 x 3-5 mm or smaller; petals 1-5-2 x as long as sepals; styles 2-2-5 x as long as 
ovary 51b. caracasanum subsp. turumiquirense(p. 90) 

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

52. el(manii(p. 90) 
Leaf lamina narrowly elliptic to linear, 0-5-2 mm wide, incurved to canaliculate; styles 
3-5-4 mm long 53. pycnophylIum{p. 92) 

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

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

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

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

elliptic to linear, narrowed at the base 55 



THE GENUS HYPERICUM L. 17 

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

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

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

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

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

l-2(-3) mm long (59. /ancioif/es) 57 

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

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

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

Flowers terminal and lateral in dense racemiform heads; habit bushy or 
straggling 59b. lancioides suhsp. congestiflorum{p. 101) 

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

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

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

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

mm long, markedly tetrastichous, glaucous 56. ilanganaticum{p. 95) 

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

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

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

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

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

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

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

styles shorter than ovary with stigmas capitate 63 

Sepal margin hyaline; inflorescence and styles not as above 72 

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

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

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

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

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

Styles 1-2 mm long; leaves dull beneath 69 

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

hyaline, dorsally usually glandular when mature 67 

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

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

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

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

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

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

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

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

38. costaricense{p . 70) 



18 NORMAN K. B. ROBSON 

Flowers 6-8 mm in diam.; leaves 3-5 mm long, subacute to obtuse; stems wiry, basally 
rooting 39. bryoides{p. 71) 

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

c.4-5 mm long 40. bolivaricuin{p. 72) 

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

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

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

42. prostratum{p. 75) 

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

marahuacanum) 73 

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

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

pseudo-dichotomous or racemiform 74 

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

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

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

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

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

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

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

tetrastichous 46. horizontale{p. 81) 

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

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

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

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

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

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

1 . Hypericum terrae-firmae Sprague & Riley 

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

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

Icon: Fig. 4. 

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

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




THE GENUS HYPERICUM L. 



19 




Fig. 4 H. terrae-firmae: (a) habit; (b) stem with leaf bases; (c) leaf scar; (d) leaf; (e) sepal; (f) petal; 
(g) stamens (partly cut away) and ovary; (h) stamen; (i) anthers; (j) capsule (a, b = I/2; c x 1 ; d, j x 2; e-g 
X 3;h X 8;i X 15). {di-d, %,]) Hunt 66; {Q,i,h,'\) Sutton etal. 193. 



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



20 





NORMAN K. B. ROBSON 

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

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

Belize (Cayo, Belize). Map 1. 

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

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



2. Hypericum styphelioides A. Richard 

in Sagra, Hist. Phys., Pol. Nat. He Cuba (Bot. - PI. Vase): 237 (1845); Leon & 
Alain, F/. Cuba3:316,t. 141 (1953);Lippoldin M55Z. Friedr. -Schiller Univ. Jena 
(Math. -Nat. R.) 19: 378, f.2. (1970). Type: Cuba, [Pinar del Rio, near San Diego 
de los Banos], Sagra s.n. (P, holotype). 

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

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

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

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



THE GENUS HYPERICUM L. 



21 





Vi 






z^ — ' 




^" 


<^^^i:^ 


: \ 






If 


'nZ::^?^ 


1 



Map 1 \. H. terrae-firmae • . 2. //. styphelioides: a. subsp. clarense A , b. subsp. styphelioides O, c. subsp. 
moaense ■. 



2a. Hypericum styphelioides subsp. clarense Lippold 

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

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

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

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

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

CUBA. Pinar del Rio: N. of San Diego de los Baiios, 13. iv. 1900 (st). Palmer & 
Riley 535 (NY, US); mountains near El Guama, 6.iii.l900 (fr). Palmer & Riley 160 
(NY, US). Las Villas: Lomas de Banao, i.l920 (fl), Lund 123 (NY); Trinidad Mts, 
Buenos Aires, near La Sabana, 5.viii.l936, Smith, Hodgson & Gonzales 3381 (F, 
NY, S, US). Isla de Pinos: Siguanea, 26.ii.1916 (fl), N. L. Britton, E. G. Britton & 
Wilson 14926 (F, NY, S, US); between Mine de Oro and Playa del Soldado, 
12.i.l956, Killip 45389 (US). 

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

2b. Hypericum styphelioides A. Richard subsp. styphelioides 

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






22 NORMAN K. B. ROBSON 

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

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

CUBA. Pinar del Rio: El Sabalo, Finca Sabanalamar, near sea level, 22.xiii.1937 
(fl), Killip 32263 (NY, US); Herradura, 1904 (fl), van Hermann 562 (BM, F, K, NY, 
P, US); near Laguna Alcatraz Grande, 16.xii.l911 (fl), Shafer 11016 (NY, US). Las 
Villas: Banao Mts., Sabana de la Gloria, 29.vii.1918 (fl), Leon & Roca 7928 (NY). 
Isla de Pinos: San Pedro and vicinity, 12.ii.-22.ii.l916 (fl), N. L. Britton, E. G. 
Britton & Wilson 14148 (F, NY, S, US); Los Indios, 4.ii.l953 (fl), Killip 42603 (NY, 
US). 

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

2c. Hypericum styphelioides subsp. moaense Lippold 

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

5/jrM6 0-6-l-5 mtall. Leaves l-\\{-\()) x 2-6 mm, obovate (or rarely oblanceolate) 
to narrowly elliptic, thickly coriaceous; slightly glaucous, ± cucullate or incurved, 
usually ± densely imbricate; apex apiculate-obtuse to acute; basal veins 5-7; glands 
prominent. Flowers 15-20 mm in diam. Sepals lanceolate, acute, 7-veined. Petals 
8-12 mm long. Styles 5(4), c. 3 mm long. 

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

Confined to Oriente. 

CUBA. Oriente: Minas de Iberia, Taco Bay, in mountains, c. 800 m, 7-8. xii. 1914 
(fl), Ekman 31K1 (F, K, MICH, NY, S); Moa, near airfield, vii.1941 (fl), Howard 
6020 (BM, NY, P, S, US); Maravi (near Barancoa), 25.xii.1914 (st), Ekman 4020 

(NY,S,US). 

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

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



THE GENUS HYPERICUM L. 

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



23 



3. Hypericum phellos Gleason 

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

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

On scrubby slopes and in moist woods immediately below them; 2600-4200 m. 

Colombia (Santander, Norte de Santander, Cesar), Venezuela (Tachira). Map 2. 

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

3a. Hypericum phellos subsp. phellos 

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

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

H. tachirense Steyerm. in sched. 

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




Q-Sa 




0-^-,_H.. 




24 



NORMAN K. B. ROBSON 




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



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

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

COLOMBIA. Norte de Santander: Paramo de Fontibon, 2600-2750 m, 15 
-16.X.1941 (fl), Cuatrecasas, Schultes & Smith 12298 (BM, COL, GH, NY, U, US); 
between Mutiscua and Pamplona, 3400 m, 23.ii.1927 (fl), Killip & Smith 19727 (GH, 
S, US); Hoya del Rio Chitaga, Quebrada de Presidente, 3100-3300 m, 28.xi.1941 
(fl), Cuatrecasas 13490 (COL, NY). Paramo de Tama, vicinity of the Cueva, 
3100-3200 m, 27.x. 1941 (fl), Cuatrecasas, Schultes & Smith 12647 (BM, F, US). 
Santander: Paramo de Santurban entre Bucaramanga & Berlin, W. side, c. 3200 m, 
3. i. 1960 (fl & fr), Barclay & Juajibioy 10422 (NY); Berlin to Picacho, 3058 m, 
14.viii.l977 (fl), Renteria, Mantilla, Nino & Ortiz 555(3) (COL, MO); vicinity of La 
Baja, 2700 m, 14-31.1.1927, Killip & Smith 18756 (GH, NY, US). 

VENEZUELA. Tachira: Paramo de Batallon, near La Grita, 3100 m, 8.iii.l979 
(fr), Kieft 104 (BM, U, VEN); SE. of Paramo de Tama, Pata de Judio, near the 
Colombia- Venezuela frontier, 2900-3000 m, 20.1.1968 (fl), Steyermark & Dunster- 
v///e 101230 (NY, VEN). 

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



THE GENUS HYPERICUM L. 25 

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

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

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

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

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

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

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

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

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

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

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

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

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

Hypericum platyphyllum Gleason in Bull. Torrey bot. Club 56: 106 (1929) pro parte, 
excl. spec. Killip & Smith 15616. Type: Colombia, Norte de Santander, Paramo de 
Romeral, 3800-4200 m, 30.i.l927 (fl), Killip & Smith 18626 (NY!, holotype; A!, 
COL,GH!,US!,isotypes). 

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



26 



NORMAN K. B. ROBSON 

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

COLOMBIA. Norte de Santander: Paramo de Santurban, extremo este, 3300 
-3500 m, 27. vi. 1940 (fl), Cuatrecasas & Garcia- Barriga 10297 (COL, F); Paramo de 
Romeral, hoya del rio Cucutilla, 3600 m, 9.iv.l973 (veg), Cuatrecasas & Jaramillo 
28751 (BM, COL, US). Santander: Paramo de Santurban, entre Berlin y Vetas, 
3700-3800 m, 22.ix.1969 (fl), Cuatrecasas & Rodriguez 27897 (BM, COL, US); 
Paramo de Romeral, 3800-4100 m, 29-30.1.1927 (fl & fr), Killip & Smith 18580 
(NY). 

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





4. Hypericum garciae Pierce 

in Bull. Torrey bot. Club 70: 174 (1943). Type: Colombia, Santander, Paramo del 
Almorzadero, Peralonso, 3200-3300 m, 20.vii.l940 (fl), Cuatrecasas & Garcia- 
Barriga 9935 (F!, holotype; NY!, US!, isotypes). 

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

Hypericum ericifolium Steyerm. in Fieldiana Bot. 28: 393 (1952) ['ericaefolium']. 
Type: Venezuela, Tachira, Paramo de Tama, 2 km above Betania, 7 km above 
Villapaez, 2500 m, 14.vii.l944 (fl), Steyermark 57212 (F!, holotype; NY!, isotype). 

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

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

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

COLOMBIA. Boyaca: Paramo de La Rusia, NW. to N. of Duitama, Serrania 
Pena Negra, Hoya de la Laguna Agua Clara, 3935 m, 10.xii.l972 (fl), Cleef 6992 
(COL, U); Vado Hondo, Siberia, between Pena de Arnical and Alto de Mogotas, 
3290 m, 4.iv.l973 (fr & fr), Cleef 9332 (BM, COL, U). Cundinamarca: Municipio de 
Tausa, paramo vecindad a la poblacion, 3200 m, 26.x. 1961 (fl), Huertas & Camargo 
5285 (COL). Meta: Paramo de Sumapaz, Hoya El Nevado, Laguna El Sorbedero, 
3550 m, l.ii.l972 (fl), C/ee/1489A (COL). Norte de Santander: Paramo de Hatico 
between Toledo and Pamplona, c. 2900 m, 12-13. iii.1927 (fl), Killip & Smith 20661 
(K, NY, US). Santander: Paramo del Almorzadero, 3600-3800 m, 28.xi.1941 (fl), 
Cuatrecasas 13496 (F, NY, P); Paramo Rico, near Vetas, 3750-3850 m, 18.1.1927 
(fl), Killip & Smith 17659 (NY). 



THE GENUS HYPERICUM L. 



27 




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



VENEZUELA. Tachira: Paramo de Tama, El Paramito, 2550 m, viii.1939 (fl), 
C/iflrrfon81(VEN). 

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

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

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

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



28 





NORMAN K. B. ROBSON 

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

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

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

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

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




6. Hypericum piriai Arechav. 

in An. Mus. nac. Montevideo 3: 108 (1898), Fl. Uruguay 1: 108 (1898); L. B. Sm. in/. 
Wash. Acad. Sci. 48: 314 (1958); Rodriguez Jimenez in Reitz, Fl. III. Catarin., 
Hipericdceas: 5, fig. 1 B (1980). Type: Uruguay, Mardonado, Pan de Aziicar cerca 
del Castillo de Piriapolis, xi.l89- (fl & fr), Arechavaleta (MVM, holotype; GH!, 
US!, photographs). 

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

H. hilaireanum L.B. Sm. in 7. Wash. Acad. Sci. 48: 314 (1958); Angely, Fl. Anal. 
Parana: 451 (1965). Type as for H. tenuifolium St.-Hil. 

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



THE GENUS HYPERICUM L. 

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

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

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

BRAZIL. Sao Paulo: Sao Paulo to Villa Emma, xii.1933 (fl), Brade 12978 (MO). 
Parana: Municipio Guarapuava, estrada para Laranjeiras do Sul, 15. xi. 1957 (fl), 
Hatschbach 4253 (US); Municipio Ponta Grossa, Parque Vila Velha, Furnas, 
i.x.l965 (fl & fr), Hatschbach 12854 (K). Santa Catarina: Municipio Abelardo Luz, 
8-12 km N. of Abelardo Luz, 900-1000 m, 15.xi.l964, Smith & Klein 13305 (NY, 
US); Municipio Xanxere, 11 km N. of Abelardo Luz, 500-600 m, 25.xii.1956, Smith 
& Klein 9231 (US). Rio Grande do Sul: Vila Manresa, prope Porto Alegre, 
2\.\.l951, Rambo 51325 (US); Porto Alegre, Morro da Policias, 21. ii. 1902 (fl & fr), 
Ma/me Regn. II 1408 (S). 

URUGUAY. Mardonado: see type. Lavalleja: Co. Penitente, 10.i.l909 (fl), 
fierro 5533 (US). 



29 





Map 4 6. H. piriai • . 



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



30 



NORMAN K. B. ROBSON 



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



? ? 







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

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

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

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

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

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

Icon: Fig. 5 A. 

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

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

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

COSTA RICA. Cartago: Volcan Irazu, 3000-3300 m, 5.V.1928 (fl), StorklOU (F, 
MICH, NY); Finca Quemado to top of Volcan Turrialba, 2800-3200 m, lO.v.1971 
(fl), Wilbur 14319 (DUKE); Cordillera de Talamanca, Cerro de la Muerte, Pan- 
American Hwy., 5 km above Millsville, 3400-3500 m, 22.vii.1949 (fl). Holm & litis 
458 (F, K, NY, P). Limon: Chirripo National Park, between Casa de Administraci6n 
and peak, c. 3400 m, 13. ii. 1983 (fl & fr), Garwood etal. 1161 (BM). San Jose: Upper 
Rio Talari, lower Valle de los Conejos and trail to Valle de los Leones, 3250-3450 m, 
21-23. viii. 1971 (fl & fr). Burger & Gomez P. 82848 (BM, DUKE, F); La Asuncion, 
Cerro Sakira, 3400 m, 27.i.l968 (fl), Wilbur & Stone 10047 (DUKE, F, MO). 

PANAMA. Chiriqui: Cerro Fabrega and vicinity near Costa Rican frontier, 
3150-3335 m, 7-8.iv.l976 (fl), Weston 10189 (MO). 

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



THE GENUS HYPERICUM L. 



31 




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



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



8. Hypericum stenopetalum Turcz. 

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



32 



NORMAN K. B. ROBSON 



} 






^5hI 


■:oy^ 




N"Y / ./['■'/ ' 


80 


75 



Map 5 1. H. irazuense A ; 8. //. stenopetalum •. 






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

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

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

Icon: Fig. 5B. 

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

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



THE GENUS HYPERICUM L. 

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

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

VENEZUELA. Merida: Paramo de Los Leones (La Lagunita, La Canada 
Grande) al W. de Murucuba, 3400-3500 m, 31. v. 1930 (fl), Gehringer 159 (F, NY, 
VEN); Paramo de San Jose, monte El Cupio, 3000 m, l.ii.l973 (fl & fr), Cuatrecasas, 
Ruiz-Teran & Lopez-Figueras 28440 (BM, US). Tachira: Paramo de Portachuelo, 
2860 m, 23.x. 1978 (fl), Luteyn 6032 (BM, MO, NY, VEN). 

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



33 



9. Hypericum carinosum R. Keller 

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

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

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

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

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

Paramo de Almorzadero, extremo sur, Peralonso, 3200 m, 19.vii.l940 (fl), 

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

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

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

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

COLOMBIA. Norte de Santander: environs of Pamplona. 10.x. 1943 (fl), de 
Garganta 626 (F). Santander: vicinity of Las Vegas, 2600-3600 m, 21-23.xii.1926, 
Killip & Smith 15921 (GH, NY, US); vicinity of California, 2800 m, 11-27.1.1927, 
Killip & Smith 16904 (F, GH, NY, US). 

VENEZUELA. Merida: Paramo de Los Leones, 3400-4500 m, 31.V.1930 (st), 
Gehringer U9{WE^). 

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





34 





NORMAN K. B. ROBSON 

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

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

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

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

Icon: Fig. 6A. 

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

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

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

COLOMBIA. Magdalena: near Rio Seville, 20.i.l959 (fl), Barclay & Juajibioy 
6557 (MO, NY); Sierra de Santa Marta, c. 48 km inland from Dibulla, c. 3850 m, 
vii.1932 (fl), Seifriz 455 (US); Sierra de Santa Marta, SE. slopes, Hoya del Rio 
Donachui, Meollaca, 3320-3260 m, 29.ix.1959, Cuatrecasas & Romero-Castaneda 
24480 (COL, US); Sierra de Santa Marta, source of Rio Donachui, 4310 m, 
22.V.1977, Starker White & Alverson 547A (NY); Sierra de Santa Marta, quebrada 
from Laguna Rio Frio, 3250 m, 28.vii.1972, Kirkbride & Forero 1751 (BM, COL). 

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



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

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



THE GENUS HYPERICUM L. 



35 




A], /ebt? 



Fig. 6 A. H. simonsii: (a) habit; (b) stem with leaves; (c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut 
away) and ovary; (g) capsule. B. H. papillosum: (h) habit; (i) leaf; (j) sepal; (k) petal; (1) stamens (partly 
cut away) and ovary; (m) young capsule (a, h x V2; b x 1 ; c-f x 4; g x 2; i-m x 3). A. Romero-Castaneda 
4541; B: (h) Grubb, Curry & Fernandez-Perez 401; (i-m) Jaramillo Mejia 212% A. 



Nevada del Cocuy, above Laguna Grande, c. 3900 m, 14.viii.l957 (fi), Grubb, Curry 
iSc Fernandez-Perez 401 (K!, holotype; US!, isotype). 

Icon: Fig. 6B. 

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



36 





-^4^ 

M 







NORMAN K. B. ROBSON 

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

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

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

COLOMBIA. Boyaca: Paramo de Pisva, road between Socha and La Punta, Km 
61, 5-5 km E. of Los Pinos, 3680 m, 14.vi.l972 (fl), Cleef 4541 (U); same road, Km 
70, 1-5 km, W. of El Cadillal, 3530 m, 16.vi.l972 (fl), Cleef 4631 (U); Paramo de 
Chita, vertiente oriental, sitio Piedras Negras, 3500-3700 m, 13.vii.l967 (fl), Jara- 
millo & van der Hammen 212% A (COL). 

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



12. Hypericum lycopodioides Triana & Planchon 

in Annls Sci. nat. (Bot.) IV, 18: 296 (1962). Type: Colombia, Cundinamarca, Andes 
de Bogota, Cipaquira [Zipaquira], viii.1855, (fl), Triana 5464 (COL?, holotype; 
BM!). 

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

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

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



THE GENUS HYPERICUM L. 



37 




Map 6 9. H. carinosum A; 12. //. lycopodioides 0;13. H. thuyoides •. 

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

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, Hoya San 
Jose, 3450 m, 9.vi.l973 (fl), Cleef 10117 (U). Boyaca: Sierra Nevada del Cocuy, 
Valle de San Jose, c. 4000 m, 10.ix.l957 (fl), Grubb, Curry & Fernandez-Perez 768 
(COL, K, US); between Chita and Sacama, Quebrada del Curial (La Porquera), 
3350 m, 15. ix. 1969 (fl), Cuatrecasas & Laureano Rodriguez 27787 (BM, COL, US). 
Cundinamarca: see type. 

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



13. Hypericum thuyoides Kunth 

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

Brathys thuyoides (Kunth) Spach, Hist. nat. veg. Phan. 5: 448 (1836), in Annls Sci. 

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

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

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

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



38 



NORMAN K. B. ROBSON 




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

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

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

COLOMBIA. Boyaca: Sogamosa to Pajarito, near Km 268, c. 2500 m, 
27.viii.1953 (fl), Langenheim 3601 (COL). Cundinamarca: Bogota to Choachi, 30 
km. Paramo de Cruz Verde, 3050 m, 8.viii.l974 (fl), Grabandt & Idrobo 219 (COL); 
Municipio de La Calera, Paramo de La Siberia, 3000-3500 m, 25.x. 1953 (fl), 
Humbert et al. 26890 (COL, P, S) Meta: Paramo de Sumapaz, Hoya Sitiales, 300 m 
an al NW. de la Laguna La Primavera, 3580 m, 25.: 72 (fl), Cleef 9S5 (COL, U). 
ToHma?: see type. 

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

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



14. Hypericum goyanesii Cuatrec. 

[in Trab. Mus. nac. Cienc. nat. Jard. bot. Madrid (Bot.) no. 27: cuadro 7 inter 54 et 55 
(1934), nomen] in op. cit., no. 29: 9 (1935), op. cit. , no. 33: 83, f. lA (1936). Type: 
Colombia, Cundinamarca, Guasca, Los Gaques wood, 3000 m, 24.iv.1932 (fl), 
Cuatrecasas 3229 (MA, holotype; K!, isotype). 

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

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



THE GENUS HYPERICUM L. 



39 



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

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

Colombia (Cundinamarca, Meta). Map 7. 



%0 





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



40 



NORMAN K. B. ROBSON 

COLOMBIA. Cundinamarca: Paramo de Guasca, 3300 m, 16.xii.l938 (fl), Balls 
5764 (BM, COL, K); Paramo de Zipaquira [Guerrero], between Zipaquira and 
Pacho, 3100-3200 m, 16.vi.l940 (fl), Cuatrecasas 9526 (COL, F, P); Paramo de Cruz 
Verde, fondo del valle al sur de La Viga, 3590 m, 28.viii.1972 (fl), Cleef 32S1 (BM, 
COL, U). Meta: Macizo de Sumapaz, airedores de la Laguna La Guitarra, 3380 
-3420 m, 4.vii.l981 (fl & fr), Diaz 2398 (COL). 

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




15. Hypericum myricariifolium Hieron. 

in Bot. Jahrb. 21: 321. (1895); R. Keller in Engler & Prantl, Nat. Pflanzenfam. 2nd 
ed. 21: 181 (1925); Gleason in Bull. Torrey bot. Club 56: 102 (1929). Type: 
Colombia, Cundinamarca, Paramo de Pasca, 3700-3800 m, vi.l868? (fl), Stubel 
Colomb. 124 (Bf, holotype; F!, NY!, photographs). 

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

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

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

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

Colombia (Cundinamarca); in the higher paramos near Bogota. Map 8 (p. 44). 
COLOMBIA. Cundinamarca: Cordillera Oriental S. of Usme, Paramo de Chisa- 
ca, c. 4120 m, 9-ll.xi.l958 (fl), Barclay & J uajibioy 6178 (COL, MO, NY); Macizo 
de Sumapaz, Andabobos, 3760-3720 m, 8. i. 1969 (fl), Cuatrecasas &Jaramillo 27031 
(BM, US); Paramo de Cruz Verde, W. slopes, 3150 m, 7.x. 1938 (fl), Cuatrecasas 409 
(COL, F, P). 

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



THE GENUS HYPERICUM L. 



41 



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

16. Hypericum cuatrecasii Gleason 

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

Shrub up to 2 m tall, spreading, with branches divergent, lateral, ± short (flowering) 
and long (extension), or pseudo-dichotomous. Stems yellow to reddish-brown, 
4-lined and ancipitous when young, eventually terete, without epidermal wrinkles or 
emergences, cortex exfoliating in strips or flakes; internodes 2-5 mm long. Leaves 
sessile, spreading to deflexed, obscurely tetrastichous, deciduous almost at the base 
after turning brown, leaving narrow basal ring; lamina (2-)4-7 x (2-)3-7 mm, 
gradually increasing in size along shoot, broadly ovate to subcircular, plane or 
slightly saccate, with midrib prominent beneath, margin not or very narrowly 
hyaline, not glaucous, coriaceous; apex acute to rounded, base cordate-amplexicaul, 
shallowly sheathing, pairs perfoliately united; basal or near-basal veins 1-3, with 
midrib branching, tertiary reticulum obscure; laminar glands dense to sparse, large, 
not prominent. Inflorescence 1-flowered, terminal and on short lateral shoots, the 
whole shoot ± cylindrical, sometimes with pseudo-dichotomous branches; pedicel 
4-7 mm long; upper leaves foliose. Flowers c. 20 mm in diam. , stellate. Sepals 5-7 x 
3-4 mm, the outer ovate, subcordiform, the inner ovate-oblong, subacute to 
rounded, cucullate, margin narrowly to broadly hyaline; veins 7-11, obscurely 
branching, midrib not prominent; glands linear, distally punctiform. Petals (bright?) 
yellow, 9-12-5 x 6-5-7-5 mm, c. 1-8 x sepals, obliquely obovate; apiculus obtuse; 
glands mostly striiform, distally punctiform. Stamens c. 120-150, longest 5-6 mm 
long,c. 0-5 x petals. Ovary 2-5-3 x l-5-2mm,broadlyovoid;styles3(4), 5-6-5 mm 
long, 2 X ovary, spreading-incurved; stigmas scarcely to narrowly capitate. Capsule 
c. 6 X 3-5 mm, ellipsoid, slightly exceeding sepals. Seeds not seen. 

In paramo and degraded forests; 1990-3000 m. 

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

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

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





17. Hypericum quitense R. Keller 

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

H. rimbachianum Diels in Biblthca bot. 116: 108 (1937). Type: Ecuador, Canar, N. 
of BibUan, c. 3400 m, 13.viii.l933, Diels 497 (Bt, holotype). 

Icon: Fig. 7. 

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



42 



NORMAN K. B. ROBSON 




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




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



THE GENUS HYPERICUM L. 



43 



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

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

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

ECUADOR. Tungurahua: around Lago Pisayambo, 3800 m, 26.xii.1964 (fl). 
Knight 268 (BM). Bolivar: Cordillera Occidental, Pucara de TeUmbela, 2600-3000 
m, 18.xi.l943 (f[),Acosta Solis 6825 (F). Caiiar: near Pimo, 3060-3120 m, 9.vii.l945 
(fl). Camp E-4133 (BM, NY). Azuay: mountains above Sayausid, 3000-3200 m, 
18.iii.l974 (fl & fr). Marling & Anderson 12630 (GB). 

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



18. Hypericum loxense Bentham 

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

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

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

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

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



^^4^ 







44 



NORMAN K. B. ROBSON 




Map 8 15. H. myricariifolium ■; 18. H. loxense: a. subsp. aequatoriale •, 
b. subsp. loxense O. 

quitense. The most primitive form of this subspecies is from Mt. Coraz6n (Cotopaxi) , 
where it is geographically isolated from the rest (Chimborazo to Loja). It is always 
erect and has incurved-cucullate, relatively broad leaves. Subsp. loxense differs in 
having relatively narrow, plane leaves and a usually laxer and more spreading habit. 
Its most primitive form occurs in Loja, whence there is (i) a southern trend (with 
disjunction) to Peru (Cajamarca, La Libertad) in which the leaves become smaller 
and narrower, and (ii) a northern trend (Azuay, Caiiar, Chimborazo) towards 
prostrate weak stems and somewhat smaller leaves. The occurrence of intermediates 
in Loja prevents the recognition of these taxa as species. 



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

H. loxense var. aequatoriale R. Keller in Bot. Jahrb. 42: 129 (1908), in Bull. Herb. 
Boissier II, 8: 181 (1908). Type: Ecuador. Canar, between Azogues and Paramo 
de Huairacaja, 2700-3000 m, ix-x.l905? (fl «fe fr), Lehmann 4795 (Bf, holotype, 
F!, photograph; F!, K!, US!, isotypes). 

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



THE GENUS HYPERICUM L. 45 

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

ECUADOR. Cotopaxi: Corazon, 3200 m, 22.vi.1876, Andre K. 436 (K). Chim- 
borazo: Paramo de Cachenco, c. 3800 m, 27.vii.1959 (fl), Barclay & Juajibioy 8258 
(NY). Caiiar: near Pimo, 3060-3120 m, 9.vii.l945 (fl), Camp E-4136 (BM, NY); 
Tipococha, 3400 m, 9.vii.l939 (fl & fr), Penland 994 (F). Azuay: Beiios, SW. of 
Cuenca, c. 2700 m, 12. v. 1974 (fl), Harling & Anderson 14561 (GB); San Miguel to 
Cuenca, 25 km, 3000 m, 15.ix.l969, B. & C. Maguire 61712 (BM, NY). Loja: 
between Loja and Cuenca, c. 20 km N. of Saraguro, 3080 m, 21.X.1980 (fl & fr), 
Croaf50854(BM,MO). 

18b. Hypericum loxense subsp. loxense 

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

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

ECUADOR. Azuay: Paramo Portete, 3600 m, 13.iii.l953 (fl), Prescottl93 (NY). 
Chimborazo: Urbina, towards Mt. Chimborazo, c. 3700 m, 27.vii.1939 (fl & fr), 
Asplund 7913 (S). Caiiar: near El Tambo (c. 69 km S. of Sibombe), 2850-3000 m, 
5.vii.l945 (fr), Camp E-4000 (BM, NY). Loja: Loja, S. of the town, c. 2350 m, 
4.X.1955 (fl & fr), Asplund 17919 (S); between Cuenca and Loja, near Una, 1865 (fl 
& fr), Jameson 127 (K, W). 

PERU. Cajamarca: above Sunchubamba, 3680 m, 8.vi.l957 (fl), Ellenberg 1862 
(U). La Libertad: Playapampa, c. 2700 m, 15-24.vi.l923 (fl), Macbride 4525 (NY). 

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

19. Hypericum sabiniforme Trev. 

Hyper. Animad. : 15 (1861); Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18: 296 
(1862) ['sabinaeforme']. Type: Colombia, Boyaca, Prov. de Tunja, Soata, 1300 m, 
iv.l843 (fl), Linden 1328 (BHU?, holotype; BM!, GH!, K!, P!, W!, isotypes). 

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

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

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

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

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

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

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

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

(1981), nomen. 

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



46 




NORMAN K. B. ROBSON 

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

In dry paramo; 2000-3870 m. 

Colombia (Boyaca). Map 9. 

COLOMBIA. Boyaca: Municipio de Susacon, paramo de Guantiva, 3140 m, 
8.viii.l958 (fl), Jaramillo & Hernandez 965 (US); NW. of Belen, Quebrada Minas, 
Hoya CUa. Larga, 3870 m, 25.ii.1972 (fl), Cleef 1807 (COL; U); 1 km E. of Vado 
Hondo, Valle de Rio Cusiara, 2915 m, 31.iii.l973 (fl). Cleef, Cuatrecasas & Jaramillo 
9236B (COL); Paramo de Giiina, 3200 m, 17.vii.l940 (fl), Cuatrecasas & Garcia 
Barriga. 9790 (F); Andes de Bogota, 2500-3000 m, (fl), Triana s.n. (BM). 

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




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



THE GENUS HYPERICUM L. 

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

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

20. Hypericum laricifoliumJuss. 

in Annls Mm. Hist. nat. Paris 3: 160, t.l6 f.l (1804); Choisy, Prodr. monogr. fam. 
Hyperic. : 58 (1821), in DC, Prodr. syst. nat. regni veg. 1: 553 (1824); R. Keller in 
Bull. Herb. Boissier 6: 257 (1898), in op. cit. II, 8: 176 (1908), in Engler & Prantl, 
Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Kunth in Reprium Spec. nov. Regni veg. 
Beih. 43: 483 (1927). Type: Peru, without precise locality or date (fl), /. deJussieu* 
in Herb. A. L. deJussieu 11800 (P- JUSS!, holotype; C!, F!, P!, isotypes). 

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

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

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

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

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

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

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

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

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

Merida, Paramo de Timotes, Paramos de Chachapos, 3800-4200 m, 24.i.l928 (fl), 

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

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

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

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



47 





48 



NORMAN K. B. ROBSON 




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



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

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

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



THE GENUS HYPERICUM L. 

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

COLOMBIA. Antioquia: between Medellin and Retiro, around Las Palmas, c. 
2800 m, 16.x. 1947 (st), Gutierrez, Barkley & Rosano 24(F). Boyaca: Paramo La 
Rusia, NNW. of Duitama, 3550 m, ll.ix.l969 (fl & fr), Cuatrecasas & Rodriguez 
27730 (BM, COL, US). Caldas: Rio Otun, Nevado de Santa Isabel, Bagas de la 
Laguna Negra, 3800-3750 m, 24.xi.1946, Cuatrecasas 23160 (F, P, U). Cauca: 
Popayan, near Pitayo, 1841-1843 (fl), Hartweg 925 (BM, K, P, W); Paramo del 
Purace, S. of the Volcan, San Francisco, 3450-3400 m, 23.vii.1943 (fl), Cuatrecasas 
14571 (COL, F, P). Cesar: Municipio de Valledupar, Corregimiento de Manaure, 
2800 m, n.d. (fl), Rodriguez s.n. (COL). Choco: Macigo de Tatama, valle El 
Encanto, 3420 m, 13. ii. 1983 (fl), Cteef & Salamanca s.n. (COL). Cundinamarca: 
Paramo de Guasca, vicinity of Bogota, 3150 m, 23.viii.1959 (fl), B. & C. Maguire 
44029 (BM, NY); Paramo de Palacio, Lagunas de Buitrago, 3550 m, 27. iv. 1973 (fl), 
Cleef9511 B (COL). Magdalena: Sierra de Perija, plain between Cerro Venado and 
Cerro Avion, 3270-3350 m, 8.xi.l959 (fl), Cuatrecasas & Romero-Castaneda 25128 
(COL) . Narino : Volcan de Cumbal , via Laguna Bolsa , 2740-2980 m , 1 1 . ix . 1944 (fl) , 
Ewan 16139 (BM, S) . Norte de Santander: Ocana to Pamplona, Serra PI. , 2400-3300 
m, 30.viii.l878 (fl), KalbreyerlW (K), Putumayo: Lagunda de la Cocha, Quebrada 
de Santa Lucia, 2850 m, 8.i.l941 (fl), Cuatrecasas 11824 (COL, F). Risaralda: 
Municipio de Pereira, vicinity of La Laguna de Otiin, 4000-4300 m, 3.ii.l980 (fl), 
Diaz, Valencia & Jaromillo 1738 (COL). Santander: Paramo de Santurban, Laguna 
de Cunta, 3880 m, 21. i. 1927 (fl), Killip & Smith 17962 (COL, F). Tolima: Paramo de 
Ruiz, 3200-3500 m, 16-17.xii.1917 (fl), Pennell 2999 (F, K, MO). Valle: Rio 



49 




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



50 NORMAN K. B. ROBSON 

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

VENEZUELA. Lara: between Buenos Aires and Paramo de las Rosas, 2285 
-3290 m, 11. ii. 1944 (fl), Steyermark 55501 (F). Merida: Paramo de Mucuchies, 
X.1865 (fl), Moritz 1166 (BM, K, P, S); Sierra Nevada de Merida, between La 
Canada del Espejo and La Canada de Los Anteojos, Torre de Befio, 4000-4130 m, 
22.1.1973 (fl), Cuatrecasas, Ruiz-Terdn & Lopez-Figueras 28570 (BM, US); Paramo 
de Conejos, c. 8 km NE. of Merida, 3800-4100 m, 27.xii.1967, Wessels Boer 2135 
(NY, U, VEN). Tachira: Paramo de La Negra, 3000 m, 13.xi.l939 (fl), Burrus & 
Miiller 1019a (VEN). Trujillo: El Paramito, Jago to La Morita, hacia Tuname, over 
3000 m, viii.1958 (fl), Aristeguieta & Medina 3488 (NY, VEN). Zulia: Zulia, 
1983-1894 (fl), Mocquerys s.n. (NY, P). 

ECUADOR. Azuay: 54 km from Cuenca on Suscal road, 3350 m, 13.ix.l969 (fl), 
B. & C. Maguire 61696 (BM, NY). Bolivar: Cordillera Occidental, Gualicon Loma, 
3240 m, l.\.\9A3,Acosta Solis 6263 (F). Canar: Biblian to Canar, north of Biblian, c. 
3400 m, 23.iv.1968 (fl). Marling, Storm & Strom 8655 (GB, NY). Carchi: Paramo El 
Angel, between El Angel and Tulcan, 3450-3500 m, 14.V.1973 (fl), Holm-Nielsen et 
al. 5298 (AAU, F, S). Chimborazo: Cerro Chiguaza, c. 3200 m, 24.ix.1968 (fl), Lugo 
475 (GB, NY). Cotopaxi: Cotopaxi, SW. slope, 3700m, 3.vii.l939 (fl), Asp lundl 516 
(K, P, S). El Oro: NE. Zaruma, Tioloma, Hac. Ambocas, 3100 m, 30.viii.l947, 
Espinosa 2034 (NY). Imbabura: Cotacache to Apuela, 21 km (Intec valley), 3300 m, 
ll.viii.l976 (fl), 0llgaard & Balslev 8687 (AAU, BM, MO, U). Loja: 10 km south of 
Loja, 2800 m, l-3.viii.1959 (fl). Marling 6239 (S). Napo: Cerro Antisana, Los 
Corrales, near Papallacta, 3900 m, 21.vii.l960 (fl), Grubb et al. 218 (K, NY). 
Pichincha: Mount Guamani, 3600-3800 m, 15.ix.l939 (fl), Asplund 8757 (K, P, S). 
Tungurahua: between Ambato and Baiios, 10. i. 1981 (fl), D'Arcy 13975 (BM, 
MO). 

PERU. Amazonas: Chachapoyas, Cordillera Calla-Calla between Leimebamba 
and Balsas, 2900-3000 m, 21.x. 1963 (fl & fr), Ferreyra & Acleto 15302 (MO, NY). 
Ancash: Cordillera Blanca, above Vicos, trail to Llacshacocha at Usacocha, 3350 m, 
14.iii.l964 (fl), Mutchinson & Wright4392 (F, K, MO, P, S). Cajamarca: 28 km N. of 
Cajamarca towards Pedernal, 12 km W. to Hacienda Granja Porcon (SIP A), 3260 
m, 6.vi.l963 (fl), D. & V. Ugent 5463 (GH, K, WIS). Huanuco: Mito, c. 2700 m, 
8-22.ii.l922 (fl & fr), Macbride & Featherstone 1610 (F, S). La Libertad: Huama- 
cucho, Pullac to Parcay, 69 km, above Rio Chuagon, 3920 m, 7.viii.l964 (fl), 
Mutchinson, Wright & Straw 6188 (F, K, MO). Piura: Huancabamba, distr. Sondor, 
2300 m, 21.vii.l975 (fl), Sagdstegui, Cabanillas & Dias 8214 (MO). San Martin: 
Huallaga, valley of Rio Apisoncho 30 km above Jucusbamba, 3300 m, 5.viii.l965 
(fl), Mamilton & Molligan 1090 (K, S). 

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

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

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

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



THE GENUS HYPERICUM L. 



51 



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

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

Icon: Fig. 8B. 

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

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

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

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

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




22. Hypericum hartwegii Bentham 

PI. Hartwegiana: 127 (1843) ['/larrweg/']; Triana & Planchon mAnnls Sci. nat. (Hot.) 
IV, 18: 295 (1862) pro parte, quoad typum; R. Keller in Engler & Prantl, Nat. 
Pflanzenfam. 3 (6): 214 (1893), in Bull. Herb. Boissier 11, 8: 177 (1908), in Engler 
& Prantl, op. cit. 2nd ed. 21: 181 (1925) pro parte, excl. pi. Colomb. Type: 
Ecuador, Loja, mountains near Loja, Chiquiribamba, vii.1841 (fl), Hartweg 721 
(K!, holotype; BM!, K!, P!, W!, isotypes). 

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



52 






NORMAN K. B. ROBSON 

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

Habitat and phenology not recorded. 

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

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






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

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

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

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

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

ECUADOR. Azuay: Cuenca to Ona, Hacienda Pizhia, c. 2800 m, 23. ix. 1955 (fl), 
Asplund 17766 (S). Azuay/Loja: Nudo del Cordillera Occidental y Cordillera 
Oriental entre Ona y Rancho Ovejero, 2704-2800 m, l-2.viii.1959 (fl), Barclay & 
Juajibioy 8452 (NY). El Oro: between Curtincapa and Guagra Uma, 13 km NE. of 
Curtincapa, 2850 m, 16.viii.l943 (fl), Steyermark 53912 (F). Loja: about Ona, 2400 
m, n.d. (fl), Jameson s.n. (K). 

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



THE GENUS HYPERICUM L. 

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

24. Hypericum magniflorum Cuatrec. 

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

Icon: Fig. 9. 

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



53 




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



54 



NORMAN K. B. ROBSON 






not deflexed, persistent until cortex is shed; lamina 10-18 x 4-8 mm, elliptic to 
oblanceolate-spathulate, plane, slightly cucullate, carinate, concolorous or paler 
beneath, not glaucous, chartaceous; apex acute to obtuse-apiculate, base angustate, 
sheathing , pairs united to form very narrow interf oliar ridge ; basal veins 5-7 , parallel 
or almost so , branching distally , with tertiary reticulation sometimes visible beneath ; 
laminar glands dense above, sparse or absent beneath, slightly prominent. Infloresc- 
ence 1 -flowered, sometimes with several short flowering branches crowded together, 
usually with pseudo-dichotomous branches from 1st or 2nd node below; pedicel up to 
5 mm long; upper leaves transitional. Flowers c. 35 mm in diam., stellate. Sepals9-\\ 
X 3-5-5 mm, broadly oblong to obovate-spathulate, acute to obtuse; veins 11-19, 
dichotomising distally, with midrib scarcely prominent beneath; glands punctiform 
in upper third. Petals rich yellow, 16-22 x 12-14 mm, c. 2 x sepals, obovate- 
oblanceolate; apiculus acute; glands punctiform distally. Stamens c. 80, longest 9-11 
mm long, c. 0-5 x petals. Ovary 4-5 x 2-3 mm, broadly eUipsoid to subglobose; 
styles 3, 5-9 mm long, 1-5-2 x ovary, erect, free, outcurved below apex; stigmas 
slightly enlarged to broadly capitate. Capsule c. 7 x 5 mm, broadly ellipsoid, shorter 
than sepals. Seeds not seen. 

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

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

COLOMBIA. Boyaca: Paramo de Chita, cabeceras de Rio Casanare, 3080 m, 
19.vi.l972 (fl), Cleef 4743 (BM, U). Norte de Santander (see also type): Bucaraman- 
ga, Cumbre of Paramo Mortino, Cucuta road, 3300 m, v. 1948 (fl), Sandeman dffll 




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



THE GENUS HYPERICUM L. 

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

VENEZUELA. Tachira: Paramo de Tama, near Colombo- Venezuelan bound- 
ary, 3045-3475 m, 15.vii.l944 (fl). Steyermark 51218 (F, NY); NNE. slopes of Pata 
de Judio, 2500-3100 m, 19.x. 1978 (fl & fr), Luteyn 5945 (BM, VEN). 

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



55 



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

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

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

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

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

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

COLOMBIA. Norte de Santander: Valley of no Chitaga, 20 km SE. of Chitaga, 
3475 m, 24.ix.1944 (fl & fr), St. John 207784 (COL, NY). Santander: Paramo del 
Almorzadero, 3600-3800 m, 28.xi.1941 (fl), Cuatrecasas 13495 (COL, GH, NY, P); 
Paramo del Almorzadero, between Chitaga and Cerrito, 3900 m, 31.xii.l959- 
l.i.l960 (fl), Barclay & J uajibioy 10389 (NY); Peralonso, Platera, 3400m, 20.ix.l969 
(fl), Cuatrecasas & Rodriguez 27828 (BM, COL, US); Peralonso, 3200 m, 
19.vii.l940 (fr), Cuatrecasas & Garcia Barriga 9887 (COL). 

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





56 



NORMAN K. B. ROBSON 

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

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





26. Hypericum mexicanum L. 

Diss. Hyperic. : 5, f . 2 {\116),Amoen. Acad. 8: 322, t.8, f .2 (1785); L. fil. , Suppl. PI. : 
345 (1781); Lam., Encyl. meth. (Bot.) 4: 169 (1797); Gleason in Bull. Torrey hot. 
Club 56: 103 (1929). Type: Colombia [Cundinamarca], Mutis in Herb. Linn. 943: 
31 (LINN!, holotype; BM!, isotype). 

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

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

Brathys mutisiana (Kunth) Spach, Hist. nat. veg. Phan. 5: 447 (1836); in Annls Sci. 
nat. (Bot.) II, 5: 366 (1836). 

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

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

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

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

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

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



THE GENUS HYPERICUM L. 



57 




Map 12 26. H. mexicanum • ; 28. //. prietoi 0;29. H. cassiopiforme I 



Paulino proximo Alto Ritacuva, c. 3650, 3. v. 1959 (fl), Barclay & Juajibioy 7525 
(COL, MO, NY); Paramo de Giiina, 3200 m, 17.vii.l940 (fl), Cuatrecasas & Garcia 
Barriga. 9783 (COL). Cundinamarca: Paramo de Guasca, 2840 m, 15.xii.l938 (fl). 
Balls 5700 (BM, COL, K); Sabana de Bogota, Municipio de Tenjo, Vereda de la 
Punta, 2800-2670 m, 29. i. 1966 (fl & fr), Cuatrecasas & Jaramillo 26755 (BM, COL, 
US); Cerro de Monserrate, 3000 m, 10.iii.l966 (fl), Duque- Jaramillo 2877 (COL). 
Meta: Macizo de Sumapaz, alredores de la Laguna La Guitarra, 3370-3400 m, 
5.vii.l981, Diaz P. 2485 (COL). Norte de Santander: Paramo de Fontibon, 2700 m, 
27.vii.1940 (fl), Cuatrecasas & Garcia Barriga. 10088 (NY, US). Santander: Paramo 
de Santurban, 3600 m, 27.viii.1948 (fl), Araque & Barkley 18s004 (COL, F). Valle: 
Rio Cali, Pichinde, 1700 m, vii.1938 (fl), Duque-Jaramillo 4107 (COL). 

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

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

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

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



58 






NORMAN K. B. ROBSON 

27. Hypericum stuebeliiHieron. 

in Bot. Jahrb. 21: 321 (1895); R. Keller in Bull. Herb. Boissier II, 8: 177 (1908), in 
Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925) Type: Peru, Amazonas, 
Chachapoyas, between Pacasmayo and Moyobamba, near Mojon Cruz, 3300 m, 
iv.-vi.l875? (fl), StubelVerull (Bt; holotype; F!, GH!, NY!, photographs). 

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

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

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

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

PERU. Amazonas: Chachapoyas, 1-5 km W. of Molinopampa, Jalca zone, 
2400-2450 m, 18.vii.l962 (fl), Wurdack 1353 (F, GH, K, NY, P, S, US); La Jalca 
near Chachapoyas, 2700-3300 m, i.l930 (fl), Williams 7585 (F, NY); Chachapoyas, 
Pass of Piscohuanuna, 2700 m, viii.1938 (fl), Sandeman s.n. (K); Cordillera of 
Piscohuanuna between Chachapoyas and Moyobamba, xii.1868 (fl), Raimondi 2096 
(NY). San Martin: Bagazan, 1835 (fl), Mathews 1609 (K). 

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

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

H. stuebelii Hieron. affinis, sed ramis strictioribus, foliis minoribus densissimis 
manifeste quadrifariis, 1-nervis, parte distali obtuse carinata parte proximaU subtus 
costa media impressa, floribus minoribus, differt. Type: Ecuador, Azuay, Eastern 
Cordillera, between Ona and the rio Yacuambi, 3000-3360 m, 10-19. ix.l945 (fr), 
Prieto P-302 (NY!, holotype). 

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



THE GENUS HYPERICUM L. 



59 



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

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

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

ECUADOR. Azuay: Eastern Cordillera between Ona and the rio Yacuambi, 
3000-3360 m, 10-19.ix.l945 (fr), Prieto P-302 (NY). 

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

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

H. stuebelii Hieron. affinis, sed ramis basalibus lateralibusque strictioribus, foliis 
minoribus, oblanceolatis vel obovato-oblanceolatis, floribus minoribus, staminis 
paucioribus, stylis brevioribus, differt. Type: Peru, Amazonas, Chachapoyas, 1-5 
km W. of Molinopampa, 2400-2450 m, 18.vii.l962 (fl), Wurdack 1355 (US!, 
holotype). 

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

In the Jalca zone; 2400-2450 m. 

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

PERU. Amazonas: 1-5 km W. of Molinopampa, 2400-2450 m, 18.vii.l962 (fl), 
Wurdack 1355 (US). 

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

30. Hypericum decandrum Turcz. 

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

Hypericum gnidioides var. polytrichoides R. Keller in Bull. Herb. Boissier II, 8: 183 
(1908). Type: Ecuador, Chimborazo, in pascuis montis Titaicun, 3900 m, xi.l859 
(fr). Spruce 5599 (G!, holotype, F!, photograph; BM!, C!, GH!, K!, NY!, P!, W!, 
isotypes). 






60 



NORMAN K. B. ROBSON 




Icon: Fig. 10. 

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

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

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

ECUADOR. Azuay: Loja to Cuenca, between Km 113 and 124, 2700-3000 m, 
26.ix.1969 (fl & fr), B. & C. Maguire 44320 (BM, NY). Caiiar: Rivera NE. of 
Pindilig, 2900 m, 13.xii.l980 (fl & fr), Holm-Nielsen, Jaramillo & Coello 29222 
(AAU). Carchi: Nudo de Boliche, Voladero, 3800 m, 12.vi.l939 (fl & fr), Penland 
900 (F, GH, NY). Chimborazo: Mt. Titaicun, 3900m, xi.l858 {^) , Spruce 5599 (BM, 
C, GH, K, NY, P, W). Loja: N. of Saraguro, 3045 m, 4.viii.l943 (fr), Steyermark 
53743 (F, NY). Morona-Santiago: Paramo de Hatanga, Km 36 on road Sigsig- 
Gualaquiza, E. of pass, 3300 m, 14.xii.l980 (fl), Holm-Nielsen, Jaramillo & Coello 
29386 (AAU). Napo: Valle de Chalupas near junction of Rio Chalupas and Rio 
Aguas Buenas, 3200 m, 20.1.1980 (fl), Holm-Nielsen 21014 (AAU). Pichincha?: 
Quito, 1865? {fi), Jameson 62 (BM, K, W). Tungurahua: Cordillera de Llanganates, 
near junction of rio Golpe and rio Sangarinas (Desaguadero), 26. xi. 1939 (fl & fr), 
Asplund 99m {S). 

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

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



31. Hypericum pimeleoides Planchon & Linden ex Triana & Planchon 

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

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



THE GENUS HYPERICUM L. 



61 




Fig. 10 H. decandrum: (a) habit, large form; (b) habit, small form; (c) stem with leaves; (d) leaf; (e) 
sepal; (f) petal; (g) stamens (partly cut away) and ovary; (h) capsule (a, b x V2; c-h x 5). (b) Espinosa 
923; others B. & C. Maguire 61708. 



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




62 



NORMAN K. B. ROBSON 




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

In open paramo; 3000-4300 m. 

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

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, Agua 
Tendida, 3100 m, 10. vi. 1973 (fl), Cleef 10174 (U). Boyaca: Sierra Nevada del Cocuy, 
near El Playon, c. 3800 m, 10. ix. 1957 (fl), Grubb, Curry & Fernandez-Perez 111 
(COL, K, US); Paramo de Chita, cabeceras de Rio Casanare, Km 93 de la carretera a 
La Punta, 3080 m, 19.vii.l972 (fl), Cleef AlAA (BM, U); Peiia de Arnical, N. de Vado 
Hondo, 3600 m, 6.iv. 1973 (fl), Cleef 9A1% (U). Cesar/Norte de Santander: 20 km al S. 




Map 13 2)1. H. pimeleoides 

wurdackii O. 



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



THE GENUS HYPERICUM L. 

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

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

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



63 



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

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

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

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

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






64 



NORMAN K. B. ROBSON 

de Rio Sevilla, 3320-3410 m, 22.i.l959 (fl & fr), Barclay & Juajibioy 6620 (COL, 
MO, NY); Sierra Nevada de Santa Marta, valley of Rio Yebosimeina, 3500-4000 m, 
22.V.1977 (fl & fr), Starker White & Alverson 547 (COL, MO, NY); Sierra de Perija, 
E. of Manaure, quebrada de Floridablanca, 2700-2800 m, 9-12.xi.l959, Cuatre- 
casas & Romero Castaneda 25171 (COL). 

VENEZUELA. Merida: Paramo del Molino, 2600 m, 19.1.1922 (fl), Jahn 940 
(VEN). Zulia: Perija, Cerro Fetari, iv.l952 (st), Urbano 2 (VEN). 

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

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




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

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

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

In paramo; 3750 m. 

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

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

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



THE GENUS HYPERICUM L. 

34. Hypericum spruce! N. Robson, sp. nov. 

H. aciculari Kunth affinis, sed caulibus crassioribus, foliis latioribus et saepissime 
longioribus, coriaceis, sepalis petalisque longioribus, staminibus numerosioribus, 
differt. Type: Ecuador, Tungurahua, [Volcan] Tungurahua, 2400-3000 m, xi.l857 
(fl), Spruce 5110 (BM!, holotype; c!, GH!, K!, P!, S!, isotypes). 

Hypericum struthiolifolium sensu R. Keller in Bull. Herb. Boissier II, 8: 180 (1908) 
Y struthiolaefoliurri']; Gleason in Bull. Torrey bot. Club 56: 102, 105 (1929), pro 
parte excl. typum. 

Icon: Fig. 11. 

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

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

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

ECUADOR. Azuay: c. 10 km NE. of Sevilla de Oro, 3000-3200 m, 13. ix. 1976 
(fl), 0llgaard & Balslev 9449 (AAU, BM, MO); 'Oriente' border. Eastern Cordil- 
lera between Ona and rio Yacuambi, 3000-3360 m, 10-19.ix.l945 (fl & fr), Prieto 
P-297 (BM, NY). Canar: between Bibhan and Canar, c. 3350 m, 21. ix. 1955 (fl), 
Asplund 17665 (S). Carchi: Base of Volcan Chiles, km34-36onTulcan-Maldonado 
road, 3900-4050 m, 19.V.1973 (fl), Holm-Nielsen etal. 5888 (AAU, F, GB, MO, S, 
U). Chimborazo: Atilio, c. 3700 m, 29.1.1968 (fl), Harling, Storm & Strom 6705 
(GB). Imbabura: Ibarra to Mariano Acosta, at pass (Hacienda Yura Cruz),' 3600 
-3800 m, 10.viii.l976 (fl & fr), 0llgaard & Balslev 8664 (AAU, BM, MO, U). Loja: 
Loja - Zamora road, 1 km east of pass (Km 12), 2725-2750 m, 17. iv. 1973 (fl), 
Holm-Nielsen et al. K. 3698 (S). Morona-Santiago: Gualaceo-General Plaza 
(Limon) road, 3000-3100 m, 20.ic.l967 (fl), Sparre 18778 (S). Napo: Cordillera de 
Los Llanganati, Chihuila Sacho o AinchiHbi, c. 3750 m, 25-29. viii. 1959, Barclay & 
Juajibioy 8996 (MO, NY); Sucumbios 'Playon de San Francisco', Camino al Cerro el 
Mirador, 3600-3800 m, 29.xii.1980 (fl), Jaramillo & Coelle 3903 (AAU, QCA). 
Pichincha: 15-4 km west of Papallacta, 3980 m, 26.iii.1972, MacBryde & Dwyer 1185 
(MO). Tungurahua: Llanganati Mts, Lake Aucacocha, 3700 m, vii.1969, Edwards 
58(K); Paramo of Minza Chica, 3800m, 4.iv.l939 (fl), Penland307 (F, GH). Zamora 
- Chinchipe: Loja to Zamora, 2800 m, 18. ix. 1961, Dodson & Thien 685 (BM). 

PERU. Piura: above Huancabamba, 3000 m, iv.l912 (fl), Weberbauer 6133 (F, 
GH, S). 

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



65 




66 



NORMAN K. B. ROBSON 




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

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

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



35. Hypericum aciculare Kunth 

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



THE GENUS HYPERICUM L. 



67 



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

H. struthiolifolium Juss. in Annls Mus. Hist. nat. Paris 3: 160 (1804) ['struthiolae- 

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

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

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

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

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

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

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

ECUADOR. Azuay: Km 85 on Pan-American Highway north of Loja, 2850-2950 
m, 3.V.1973 (fl), Holm-Nielsen et al. 4808 (AAU, F, GB, S, U); Rio Collay south of 
El Pan, Huagrarancha slopes, 2650-3290 m, 6.vii.l943 (fl), Steyermark 53372 (F); 
*30-50 km S. of Cuenca, Paramo de Tinajillas; 3300-3450, 17.iii.l945, Camp 
E-2229 (BM, NY);* Rancho Ovejero, c. 3000 m, 1-2. viii. 1969 (fl). Barclay & 
Juajibioy 8484 (NY). Loja: between La Toma and Loja, 1800-2600 m, 4.ix.l923 (fl), 
Hitchcock 21431 (GH, NY, US);* 7 km from Loja on road to Zamora, 2350 m, 
8.ix.l959 (fl), B. & C. Maguire 61667 (BM, NY);* W. slopes of Cerro Villonaco, 
2200-2250 m, 10.iv.l974 (fl & fr), Harling & Anderson 13351 (GB, NY). 

PERU. Piura: Huancabamba to Cuello del Indio, Mitopampa, 2650 m, 22.vii. 1975 
(fl), Sagdstegui, Cabanillas & Dios 8253 (MO). Amazonas:* Chachapoyas, 3-6 Km 
W. of Molinopampa, 2200-2450 m, 19.vii.l962 (fl), Wurdack 1403 (F, K, NY, S, 
US). *Pajonales, Pozuro, 3000 m, vii.1863 (fl), Pearce 252 (BM, K). 

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



l\\ 



W 




Wet habitat form, see p. 68. 



68 



NORMAN K. B. ROBSON 




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

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

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



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

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

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



THE GENUS HYPERICUM L. 



69 



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

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

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

PERU. Amazonas: Chachapoyas, Cordillera Calla-Calla between Leimebamba 
and Balsas, 2900-3000 m, 21.viii.l963 (fl), Ferreyra & Acleto 15284 (NY); between 
Leimebamba and Balsas, 2700 m, l.vi.l963 (fl & fr), Lopez, Sagdstegui & Callantes 
4429 (NY); Cerro Calla-Calla, E. side, 19 km above Leimebamba on Balsas road, 
3100 m, 4.vi.l964 (fl), Hutchinson & Wright 5505 (F, NY, UC, US); Jalca de 
Calla-Calla, 3100 m, 7.V.1970 (fl & fr), Sagdstegui 7451 (F, NY, VEN); Chachapoyas 
to Moyobamba, La Jalca, 2700-3300 m, 20.i.l930 (fl), Williams 7583 (F, NY); 
Chachapoyas, 1826 (fl & fr), Mathews 2130 (BM, K, P). Junin: Huancayo, Quebrada 
Occopilla, ii.l948 (fl), Soukoup 3636 (MO). Pasco: Oxapampa, Cordillera Yana- 
chaga, Cerro Pajonal 12 km SE. of Oxapampa, 2700-2800 m, 7.x. 1982 (fr), Foster 
8992 (BM, MO). 

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

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

H. recurvo N. Robson affinis, sed foliis longioribus, carinatis, densissime imbricatis, 
valde tetrastichis alas formantibus, floribus maioribus, differt; a H. aciculare Kunth 
isdem characteribus differt. Type: Peru, Amazonas, Chachapoyas, summit of Cerro 
Malcabal (Cerro Tumbe), 3-6 km SW. of Mohnopampa, 2850-2900 m, 20.vii.l962 
(fl & fr), Wurdack 1426 (K!, holotype; F!, GH!, MICH!, NY!, P!, S!, US, isotypes). 

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







70 



NORMAN K. B. ROBSON 



No habitat data known; 2850-2900 m. 

Peru (Amazonas-Chachapoyas). Known only from type collection. Map 13 (p. 62). 
PERU. Amazonas: Chachapoyas, summit of CerroMalcabal(CerroTumbe), 3-6 
km SW. of Molinopampa, 2850-2900 m, 20.vii.l962 (fl & fr), Wurdack 1426 (F, GH, 
K,MICH,NY,P,S,US). 

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




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

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

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

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

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

COSTA RICA. Alajuela: upper slopes of Volcan Poas, c. 2700 m, l.ix.l968 (fl «fe 
fr), Wilbur & Stone 10634 (BM, DUKE, GH, MO, NY, US). Cartago: Asuncion of 
Cerro de la Muerte, c. 3150 m, 17.V.1971 (fl), Wilbur 14563 (DUKE, MICH, MO). 
Limon: Chirripo National Park, between Casa da Administracion and peak, c. 3400 
m, 13. ii. 1983 (fl & fr), Garwood et al. 1162 (BM). Puntarenas/Bocas del Toro 
(Panama): Cordillera de Talamanca, Cerro Echandi, on the International border, 
3050-3160 m, 22.viii.1983 (fl), Davidse etal. 2389 (BM, MO). 

PANAMA. Bocas del Toro: Valle de Silencia, 2440 m, 8.viii.l979 (fl & fr), 
Antonio 1547 (BM, MO). Chiriqui: Cerro Fabrega and vicinity, near Costa Rican 
frontier, 3150-3335 m, 7-8.iv.l976 (fl), Weston 10149 (MO). 

COLOMBIA. Antioquia: Paramo Frontino, near Llano Grande, 3450 m, 
25.x. 1976 (fl & fr), Boeke & McElroy 229 (NY), Bohvar/ Antioquia: Paramo de 
Chaquiro, 900-960 m, 23.ii.1919 (fl & fr), Pennell 4285 (GH, NY). 



THE GENUS HYPERICUM L. 



71 







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



I,b. 



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

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



39. Hypericum bryoides Gleason 

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

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



ylTW 




72 



NORMAN K. B. ROBSON 



9 P 




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

Rocky places in open paramo; 3000-4200 m. 

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

COLOMBIA. Boyaca: Sierra Nevada del Cocuy, Alto La Cueva, 3850 m, 
5.vi.l973, Cleef 10000 (BM, COL, U). Norte de Santander: Paramo de Romeral, 
3800-4200 m, 30.1.1927 (st), Killip & Smith 18631 (F, GH, K, NY, S, US); Paramo 
de Santurban, 5 km E. of Berlin, 3200 m, 4.vii.l984 (fl), 1^00^4504 (K). Santander: 
Paramo de Almorzadero, 3500-3700 m, 20.vii.l940 (fl), Cuatrecasas & Garcia 
Barriga 9993 (F), 9995 (COL), 9996 (NY). 

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

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

H. magdalenico N. Robson affinis, sed foliis angustissime ellipticis basi angustatis 
saepe terminantibus, petalis brevioribus, staminibus paucioribus, stylis brevioribus 
stigmatibus manifeste capitatis, differt. Type: Colombia, Bolivar, headwaters of Rio 
Sinu, 1918 (fl), Pennell 4783 (NY!, holotype). 

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

No habitat or altitude cited. 

Colombia (Bolivar). Known from only one collection. Map 16 (p. 74). 
COLOMBIA. Bolivar: headwaters of Rio Sinu, 1918 (fl), Pennell 47S3 (NY). 

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

41. Hypericum juniperinum Kunth 

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



THE GENUS HYPERICUM L. 

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

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

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

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

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

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

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

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

H. strictum var. reductum Gleason in Bull. Torrey bot. Club 56: 104 (1929). Type: 
Colombia, Santander, west slope of the Paramo de San turban towards Tona, 3000 
m, 18.ii.l927 (fl & fr), Killip & Smith 19534 (NY!, holotype). 



73 



UJ 



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



9 o 



^\^ 

^"y" 



74 



NORMAN K. B. ROBSON 




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

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

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

COLOMBIA. Antioquia: Paramo de Urrao, 3500 m, ix. 1960 (fl), Espinal & Perez 
258 (COL); 1 km norte de Santa Rosa de Osos, c. 2600 m, 25.ix.1948 (fl & fr), Perez 
& Barkley 18A058 (COL, NY); Paramo Frontino, near Llano Grande, 3450 m, 
25.V.1976 (fl & fr), Boeke & McElroy 219 (MO, NY). Boyaca:* between Soata and 
Cocuy, Paramo del Alto del Canutal, 3300-3400 m, 8.ix.l938 (fl & fr), Cuatrecasas 
1197 (F); Sierra Nevada del Cocuy, between Praton and Cobugon, c. 3100 m, 
15.viii.l957 (fl), Grubb, Curry & Fernandez-Perez 507 (COL, K); Guican, Hoya del 
rio Tabor, en direccione a Ritacuva, 3570-3530 m, 18. ix. 1969 (fl), Cuatrecasas & 
Rodriguez 27833 (BM, COL, US); Paramo de la Rusia, NW.-N. de Duitama, cerca 
del puente de rio Surba, 3240 m, 9.xii.l972 (fl & fr), Cleef 6925 (COL, U); carreteria 
Sogamoso-Pajarito, Km. 57-73, 2850-3010 m, 20.ix.l969 (fl), Diaz 196 (COL). 
Caldas: cabeceras del rio Otun, hacia el Nevado de Santa Isabel, Bagas de la Laguna 
Negra (Quebrada de la Leona), 3800-3750 m, 24.xi.1946 (fl), Cuatrecasas 23159 
(BM, F, P, U). Cuaca: Valley of Rio Cofre, near Gabriel Lopez, c. 3100 m, 14.x. 1961 
(fl & fr), Cuatrecasas & Willand 26431 (COL); Paramo de Guanaco, 1843 (fl), 
Hartweg 926 (BM, K, P, W);* Macizo Colombiano, Valle de Las Papas, alredores de 
Valencia, Los Andes, margens del ri Sucubun, 2910 m, 11. ix. -1.x. 1958, Idrobo, 
Pinto & Bischler 3725 (COL, P). Cundinamarca: Macizo de Bogota, El Retire, 
Bogota, 2600-2700 m, (v.l946 (fl & fr), Schultes 7004 a (COL, F, GH); Bogota to 
Choachi, 27 km W. of Choachi, 3260 m, 4.i.l976 (fl), Luteyn 4730 (BM, COL, MO, 
NY); Paramo de Guasca, 2840 m, 15.xii.l938 (fl), Balls 5680 (BM, COL, F, K); 
Paramo de Sibate, 3570 m, 22.xi.1981 (fr), Inglesias 19 (COL); entre Bogota y La 
Calera, 2650-3000 m, 27.xi.1947 (fl), Barkley, Garcia- Barriga & Vanegas 17C814 
(COL, US). Magdalena: Sierra Perija, near Laguna el Juncos, above 'Africa', c. 
2200 m, 16.xii.l944 (fl & fr), Haught 4503 (F, K). Meta: Paramo de Sumapaz, Hoya 
El Nevado, Laguna La Guitarra, 3460 m, 21.i.l972 (fr), Cleef S24 (COL, U). Norte 
de Santander: Paramo del Hatico (between Toledo and Pamplona), c. 2900 m, 
12-13.iii.1927 (fl), Killip & Smith 20662 (GH, NY). Putumayo: between Laguna La 
Cocha and Paramo de Tabano, c. 2800-3000 m, l.i.l946 (fl), Schultes & Villareal 
7834a (F). Santander:* Bucaramanga (Cumbre to Cucuta), 3300 m, v.1948 (fl), 

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



THE GENUS HYPERICUM L. 



75 



Sandeman 6067 (BM, K); Paramo de Almorzadero, 3600-3800 m, 28.xi.1941 (fl), 
Cuatrecasas 13493 (COL, F, NY); limites entre los Departamentos Santander y 
Boyaca, Corregimiento de Virolin, Finca 'La Sierra', 2500-2600 m, 18. vi. 1976 (fl), 
Lozano, Torres & Diaz 2636 (COL). Tolima: 'Rosalito', near Paramo de Ruiz, 
2800-3100 m, 15-17.xii.1917 (fl & fr), Pennell 2956 (F, GH, K, MO, NY). 

VENEZUELA. Merida: Paramo de La Negra above Bailadores, 3100 m, 
13. ii. 1939 {^), Alston 7024 (BM, S, U);* Paramo de Muruti, x.1865 (fl), Moritz 1165 
(BM, GH, K, P, S) ; Paramo La Negra between Tovar and La Grita, 3000 m, 7.x. 1965 
(fl), Breteler A(>\9 (K, MO, NY, S, U, VEN). Tachira: Paramo Zumbador, 14 km S. 
of El Cobre, 2500 m, 31.iii.l974 (fl). Gentry, Morillo & Morillo 11068 (GH, MO, 
NY, VEN); Dtto. Uribante, a las cabeceras del rio Uribante, 51 km al SW. de 
Pregonero, 2700-2900 m, 28. ix. 1981 (fl), Steyermark & Manara 125420 (BM, VEN). 
Trujillo: Alredores de Guirigay, hacia Laguna La Parida, 3300 m, viii.1958 (fl), 
Aristeguieta & Medina 3537 (NY, VEN). 

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

In Cauca there is a separate apparently continuous morphocline in which the styles 
are always three in number. This cline links the tall erect plants with crowded 
subappressed leaves and larger flowers (i.e. H. juniperinum sensu Spach and Triana 
& Planchon) with smaller, spreading to decumbent plants with stems rooting, longer 
internodes, smaller, closely appressed leaves, and smaller flowers. In its extreme 
expression, the latter form approaches H. prostratum, but can be distinguished from 
it by the larger leaves and flowers and the longer styles. Specimens exemplifying this 
end of the morphocHne include Idrobo, Pinto & Bischler 3725 (COL, P), 3717 (COL) 
and Barclay & Juajibioy 5784 (COL, MO, NY), 5809 (COL, MO, NY), all from 
Valle de Los Papas, Los Andes, near Valencia. 



42. Hypericum prostratum Cuatrec. 

in Brittonia 11: 166 (1959). Type: Colombia, Cundinamarca, Paramo de Sumapaz, 
3800 m, 10.vi.l952 (fl), K0ie 4621 (US!, holotype; C!, isotype). 

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

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






76 



NORMAN K. B. ROBSON 

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

COLOMBIA. Boyaca: Paramo de Pisva, carretera Socha-La Punta, Km 61.5, 6 
km al E. de Los Pinos, Alto de Granados, 3570 m, 11. vi. 1972 (fl), Cleef 4374 (BM, 
COL, U); Paramo de La Rusia, NW.-N. de Duitama, Aislada, 3575 m, 6.xii.l972 (fl 
& fr), Cleef 6747 (COL, U); between Soata and Cocuy, Paramo del Alto del Escobal, 
3750 m, 8.ix.l938 (fl), Cuatrecasas 1224 (F). Cundinamarca: Bogota-Usme road. 
Paramo de Chisaca, between Kms 28 and 33, 3375-3450 m, 17.V.1979 (fl), Luteyn 
7770 (BM, COL, MO, NY); Paramo de Cruz Verde, aprox. 5 km al ENE. de la 
Laguna El Verjon, 3360 m, 2.V.1972 (fl), Cleef 3371 (BM, COL, U); entre Cogua y 
San Cayetano, Laguna Verde y alredores, 3600 m, 12.xi.l972 (fl), Cleef 6160 (BM, 
COL, U). Meta: Paramo de Sumapaz, Hoya El Nevado, Laguna La Guitarra y 
alredores, 3460 m, 21. i. 1972 (fl), C/ee/834 (BM, COL, U). Santander: Paramo de 
Almorzadero, 3500-3700 m, 20.vii.l940 (fl), Cuatrecasas & Garcia Barriga 9998 
(COL, US). 

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



^^ 



¥ 




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

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

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

Paramo? ; 3700-3960 m. 

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

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

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



THE GENUS HYPERICUM L. 



77 




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



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

H. magdalenico N. Robson affinis, sed foliis minoribus acicularibus minus coriaceis, 
costa media baud ramosa, sepalis angustioribus stylis gracilioribus brevioribus, inter 
alia differt; a H. lancifolio Gleason foliis angustioribus lucentibus torquescentibus 
margine valde incurvatis differt. Type: Venezuela, Amazonas, Dept. Atabapo, 
Cerro Marahuaca, cumbre Seccion suroriental, 2685 m, 15.1.1981, Maguire, Steyer- 
marketal. 65629 (VEN!, holotype; MO!, NY!, isotypes). 

Icones: Fig. 12A-C. 

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

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

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



lii 



o o 



y V 



a,c 





78 



NORMAN K. B. ROBSON 




Fig. 12 H. marahuacanum. A. subsp. marahuacanum: (a) habit; (b, b') stem with leaves (two aspects); 
(c) leaf; (d) sepal; (e) petal; (f) stamens (partly cut away) and ovary; (g) capsule. B. subsp. strictissimum: 
(h) habit. C. subsp. chimantaicum: (i) habit (a, h, i X V2; b-g X 3). A. Maguire etal. 65629; B. Fosberg 
22209; C. Huber & Steyermark 7014. 



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

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



THE GENUS HYPERICUM L. 79 

44a. Hypericum marahuacanum subsp. marahuacanum 

Icon: Fig. 12A. 

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

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

Venezuela ( Amazonas) . 

VENEZUELA. Amazonas: Departamento Atabapo, Cerro de Marahuaca, parte 
central de la Meseta S.-E., Quebrada Yekuana, afluente del rio Negro, 2560 m, 
10-12.X.1983, Steyermark 129511 (BM, VEN); Cerro de Marahuaca, al NE. de, y 
casi contiguacon, Cerro Duida, c. 2750 m, 2 & 9.ii.l975, Tillett, Colveeetal. 752-335 
(VEN). 

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

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

a subsp. marahuacano ramulis plerumque lateralibus, foliis angustioribus, floribus 
minoribus, sepals 3-5-venatis, petalis quam sepalis 1 •7plo longioribus, stylis 2-5-3 
mm longis, inter alia differt. Type: Colombia, Boyaca, Socha, valley of Rio 
Chicamocha, 2700-2750 m, 8.xi.l944 (fl), Fosberg22209 (US!, holotype; NY!, P!, 
isotypes). 

Icon: Fig. 12B. 

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

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

Colombia (Boyaca), Venezuela (Merida). 

COLOMBIA. Boyaca: Carretera Socha-Los Pinos, Km 41, 2 km al N. del Alto 
Los Pinos y 4 km al ENE. de Socha, 2970 m, 22.V.1973 (fl), Cleef 9891 (COL, U); 
Paramo de Belen, c. 14 km N. of Belen, near 259 km marker (from Bogota), c. 3250 
m, 9.V.1959 (fl), Barclay & Juajibioy 7668 (NY). 

VENEZUELA. Merida: Paramo de Las Coloradas (entre Santa Cruz de Mora y 
El Molino), San Rafael, 2950-2950 m, 9.ii.l973 (fl), Cuatrecasas, Ruiz-Terdn & 
Lopez-Figueiras 28529 (BM, US); Sierra Nevada, i.l865 (fl), Moritz 1602 (BM). 

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



80 NORMAN K. B. ROBSON 

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

44c. Hypericum marahuacanum subsp. chimantaicum N. Robson, subsp. 
nov. 

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

Icon: Fig. 12C. 

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

In swampy open savanna; 2121-2200 m. 

Venezuela (Bolivar: Macizo de Chimanta and Roraima). 

VENEZUELA, Bolivar: Distrito Piar, Macizo de Chimanta, E., Apacara-tepui, 
sector Norte de Macizo, c. 2200 m, 30.i-l.ii.l983 (fl), Steyermark, Huber & Carreno 
128351 (BM, VEN), Huber & Steyermark 7014 (BM, VEN); Cimo del Roraima, 
i.l977 (fl), Delascio & Brewer 4794 (VEN). 

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

45 . Hypericum lancifolium Gleason 

in Bull. Torrey bot. Club 56: 103 (clav.), 105 (1929). Type: Colombia, Santander, 
vicinity of La Baja, 2200-2600 m, 14.i.l928 (fl), Killip & Smith 17138 (NY!, 
holotype; US!, isotype). 

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

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

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

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



THE GENUS HYPERICUM L. 



81 



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

In paramo; c. 2600-3700 m. 

Colombia (Boyaca, Santander, N. de Santander), Venezuela (T^chira, Apure). 
Map 17 (p. 77). 

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

VENEZUELA. Apure: Paramo de Pata de Judio, en la frontera Colombo- 
Venezolana, 30 km al sur de San Vicente de la Revancha, SE. del Paramo de Tama, 
19.1.1968, Steyermark & Dunsterville 10122 (NY, VEN). Tachira: Paramo de Tama, 
cerca de la frontera Colombo- Venezolana, 3000-3200 m, 20-23. v. 1967, Steyermark 
& Dunsterville 98752 (VEN). 

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

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

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



9 9 9 9 




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

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

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




82 



NORMAN K. B. ROBSON 



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

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

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

COLOMBIA. Norte de Santander: Paramo de Santurban, en route from Tona to 
Mutiscua, 4200-4500 m, 19. ii. 1927 (fl & fr), Killip & Smith 19564 (BM, GH). 
Santander: Paramo de Las Vetas, a 2 km del paraje de Berlin, c. 2800 m, 18.xii.l948 
(fl & fr), Molina & Barkley 18S5410 (COL, F); idem, c. 3450 m, 3.vi.l960, Barclay & 
Juajibioy 10445 (MO, NY); Paramo de Almorzadero, 4200 m, 9.ix.l955 (fl), 
Vareschi 4070 (VEN p.p.); idem, auf der Passhohe, 3850 m, 24.vii.1952, Schneider 
1232 (S). 

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



k 



Al . Hypericum tetrastichum Cuatrec. 

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

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




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



THE GENUS HYPERICUM L. 

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

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

Colombia (Boyaca, Santander, Venezuela (Merida). Map 18. 

COLOMBIA. Boyaca: Municipio Villa de Leyva, Parque Nacional Santuario de 
Iguaque, 3700 m, 20.vii.l979, Melampy 161 (COL); Municipio de Duitama, Paramo 
de La Rusia, 3400-3500 m, 28.xi.1978 (fl), Diaz P. 1243 (COL); Paramo de 
Guantiva, 3100 m, 26.vi.1964, Espinal 1733 (COL); paramos al NW. de Belen, 
vereda S. Jose de la Montana, Alto de las Cruces y alredores, Cabeceras Q. El Toral, 
c. 3950 m, 24.ii.1972 (fl), C/ee/1736 (COL, U). Santander: Paramo Ruso, 3150 m, 
12.vii.l968 (fl), Barclay & Mullen 38C137 (COL). 

VENEZUELA. Merida: Sierra Nevada de Merida, margen N. de la Laguna verde 
(al pie del Pico Humboldt), ± 4100 m, fl. 15-18.ii.l966, Schulz, Rodriguez & 
Ramirez 35S(\J). 

H. tetrastichum is closely related to H. lancifolium and H. strictum, differing 
essentially from the latter in its markedly tetrastichous foliage and in the length of the 
styles. The leaves appear pungent but may not be rigid when alive. 

In some Boyaca locahties, H. tetrastichum occurs in two apparently distinct forms: 
(i) leaves and flowers larger; leaves glandless beneath, apex acuminate-acicular 
(damp habitats); (ii) leaves and flowers smaller; leaves glandular beneath, apex 
acute (dry habitats). Until more information is available on the nature of this 
variation (genetical or environmental), it seems preferable not to give form (ii) any 
taxonomic recognition. Specimens of this form include: 

Paramo de La Rusia, NW.-N. de Duitama, Laguna Negra, 3725 m, 14.xii.l972 
(fl), CleefllOS (COL, U), 7208A (COL); Paramos al NW. de Belen, cabeceras 
Quebrada Minas, Hoya Laguna El Alcohol, 3900 m, 26.ii.1972 (fl), Cleef 1S44 (U). 

The drawing of the ovary accompanying the description of H. tetrastichum by 
Cuatrecasas depicts the styles as shorter than the ovary. However, my observations 
indicate that they are as long as or longer than the ovary, and suggest that the ovary 
depicted by Cuatrecasas had begun to enlarge after fertilization. 



83 



go q g 





48. Hypericum strictum Kunth 

in Humboldt, Bonpland & Kunth, Nova Gen. et Sp. PI. 5: 190 (1822); Gleason in 
Bull. Torrey hot. Club 56: 104 (1929) pro parte excl. vars. Type: Columbia, 
Cundinamarca, prope Santa Fe de Bogota, 2880 m, 1805 (fl), Humboldt & 
Bonpland (P!, holotype; US!, photograph). 

Shrub 0-2-1 m tall, erect, with branches strict, pseudo-dichotomous or mostly 



84 



NORMAN K. B. ROBSON 




lateral, sometimes decumbent and rooting at the base. Stems orange-brown, 4- 
angled and ancipitous when young, then 2-lined, subquadrangular, eventually 
terete, cortex exfoliating in strips; internodes 1-7 mm long. Leaves sessile, imbri- 
cate, outcurving or spreading but not twisting, not or obscurely tetrastichous, 
deciduous above base without fading; lamina 5-13 x 5-6-1-3 mm, linear-acicular, 
incurved-canaliculate, cucuUate or not, midrib impressed dorsally, margin narrowly 
hyaline, sometimes red-tipped when young, otherwise concolorous, dull to sub- 
lucent on both sides, glaucous when young, coriaceous to chartaceous; apex acute to 
subacuminate, base parallel-sided, pairs forming narrow interfoliar ridge; basal vein 
1, unbranched; laminar glands absent or rarely sparse and visible beneath. Inflores- 
cence 1-flowered, terminal and sometimes on short to long lateral branches from up 
to 5 nodes below (racemiform), sometimes with pseudo-dichotomous branches from 
node below; pedicel 2-5 mm long, rather slender to rather stout, not incrassate 
upwards; upper leaves not transitional. Flowers c. 12-20 mm in diam., stellate to 
obconic. Sepals 4-5-10 x 1-2 mm, lanceolate to narrowly oblong, acute to acutely 
acuminate, veins 3-5, unbranched, not or scarcely prominent but glands often ± 
depressed ('sepals ribbed'); margin ± broadly hyaline; glands linear, distally inter- 
rupted. Petals bright yellow, 6-11 x 3-7 mm, 1-1-1-3 x sepals, oblong-obovate to 
obovate; apiculus acute to apiculate; glands interrupted, distally subpunctiform. 
Stamens 35-50, longest 4-7 mm long, c. 0-65 x petals. Ovary 1-3-2 x 1-1-5 mm, 
ellipsoid to ovoid-ellipsoid or subglobose; styles 3, 2-3 mm long, 1-1-5 x ovary, 
ascending; stigmas narrowly to broadly capitate. Capsule 4-6 x 2-5-3-5 mm, 
ellipsoid, acute to obtuse, shorter than sepals. Seeds 0-6-0-8 mm long, ecarinate; 
testa finely scalariform-reticulate. 

In exposed areas in paramo and subparamo, and open grassland; 2120-4000 m. 

Colombia (Santander, Boyaca, Cundinamarca, Meta). Map 19. 

H. strictum is closely related to H. tetrastichum, differing from it in the leaves, which 
soon cease to be clearly tetrastichous as a result of stem internode elongation and leaf 
outcurving. In addition the leaf apex is never so sharply acuminate as it is in the 
longer-leaved form of//, tetrastichum, and the sepal form and glands are distinct. H. 
strictum also has a different, though overlapping distribution, being centred in 
Cundinamarca and showing trends north-east and south-west from there along the 
Cordillera Oriental. Some Boyaca specimens verge toward H. tetrastichum morpho- 
logically; others are quite distinct from that species and, indeed, form a derivative 
subspecies of //. strictum. 



48a. H. strictum subsp. strictum 

H. struthiolifolium var. strictum (Kunth) Choisy in DC. , Prodr. syst. nat. regni veg. 1: 
553 (1824) ['struthiolaefolium']; Weddell, Chloris Andina 2: 271 (1857); Triana & 
Planchon mAnnls Sci. nat. (Bot.) IV, 18: 294 (1862). 

Leaves 8-13 mm long, outcurving, with apex subacuminate to acute, not cucullate, 
rarely red-tipped when young, dull, usually eglandular beneath. Stems simple, 
usually erect. Sepals 5-10 x 1-3-2 mm, lanceolate, acutely acuminate to acute, 
usually outcurving, rarely red-tipped; veins 5; glands adjacent to midrib often 
depressed. Petals 8-11 mm long. Stamens c. 50, longest 4-7 mm long. Ovary 1-5-2 
mm long, ellipsoid to ovoid-ellipsoid. Seeds 0-7-0-8 mm long. 

In dry rocky paramo and open grassland. 

Colombia (Boyaca, Cundinamarca, Meta). 

COLOMBIA. Boyaca: Rio Grande, 2 km N. of Combita, 3445 m, l.ix.l944 (fl), 
St. John 20695 (NY, US); NW. of Belen, Hoya El Pulpito, 2 km ESE. of Laguna 
Grande, 3805 m, 6.V.1973 (fl), Cleef 9775 B (COL). Cundinamarca: Represa del 
Neusa, 3350 m, 22.vi.1957 (fi), Barclay 4157 (COL); Zipaquira - Pacho, Paramo 
Alto, ll.iii.l951 (fl), Romero-Castaheda 2282 (COL, F); Paramo de La Calera, 
3000-3300 m, 25.ii.1950 (fl & fr), Philipson, Idrobo & Fernandez 2440 (BM, COL, 
F); Bogota, paramo above El Chico, 3150-3300 m, ll.vii.l943 (fl), Fosberg & 



THE GENUS HYPERICUM L. 



85 




Map 19 48. H. strictum: a. subsp. strictum ■ , b. subsp. compactum ('gracile' form) O, ('compactum' 
form) •. 

Villareal 20585 (COL, NY, US). Meta: Paramo de Sumapaz, Cerro Nevado del 
Sumapaz, 3720 m, 31.i.l972 (fl), Cleef 1535 (U). 

H. strictum subsp. strictum can be recognized by its dull, outcurving, usually 
apparently eglandular leaves and the lanceolate sepals that are not or scarcely ribbed 
but often have the two linear glands on either side of the midrib impressed. 



48b. H. strictum subsp. compactum (Triana & Planchon) N. Robson, 
comb, et stat. nov. 

H. pseudobrathys Turcz. in Bull. Soc. Nat. Moscou 31 (1): 387 (1858) ['pseudo- 

brathys'] pro parte, quoad spec. Funck & Schlim 1279, Linden 1313. 
H. struthiolifolium [var.] y compactum Triana & Planchon [in Weddell, Chloris 

Andina 2: 271 (1857) ['struthiolaefolium\ 'confertum'], nomen] in Annls Sci. nat. 

(Bot.) IV, 18: 294 (1862). Type: Colombia, Santander, prov. de Pamplona, San 

Urban, 2570 m, 1846 (fl), Funck & Schlim 1279 (COL?, holotype; BM!, P!, 

isotypes). 
H. brathys sensu Trev., Hyper. Animad.: 15 (1861) pro parte, quoad spec. Linden 

1313. 
H. struthiolifolium [var.] e grad/e Triana & Planchon [in Weddell, Chloris Andina 2: 

271 (1857), nomen] in Annls Sci. nat. (Bot.) IV, 18: 294. (1862), {struthiolae- 

folium"] nom. illegit., pro parte, quoad spec. Linden 1313. 
H. laricifolium var. glaucum R. Keller in Bull. Herb. Boissierll, 8: 182 (1908). Type: 

Colombia, Boyaca, prov. Tunja, Paramo de Coati, pres de Serinsa, 2870 m, 

1842-1843 (fl), Linden 1313 (W!, holotype; GH!, KW!, P!, W!, isotypes). 
H. strictum var. compactum (Triana & Planchon) Gleason in Bull. Torrey bot. Club 

56:104(1929). 
H. graciliforme N. Robson in Cleef, Veg. Pdram. Colomb. Cord. Or.: 304 (1981), 

nomen. 

Stems usually branched from the base, ± ascending. Leaves 5-7(-8) mm long, 
outcurving or spreading with apex subacute, ± cucullate, markedly red-tipped or 
-tinged when young, dull to sublucent, often glandular beneath. Sepals 4-5-5-5(-6) 
X l-l-2(-l-5) mm, triangular-lanceolate to narrowly oblong, acute to subacute, not 
outcurving, red-tipped or -tinged; veins 3(-5), all ± impressed. Petals 6-8(-10) m 



\D 



86 





NORMAN K. B. ROBSON 

long. Stamens 30-40, longest 4-5 mm long. Ovary 1-3-2 mm long, ellipsoid- 
subglobose to subglobose. Seeds 0-6-0-7 mm long. 

Wet areas in paramo and subparamo, often with Espeletia spp. 

Colombia (Boyaca, Santander). 

COLOMBIA. Boyaca: Paramo de Santa Rosa, entre Santa Rosa de Viterbo y 
Cerinza, 2800 m, 17.vii.l940 (fr), Cuatrecasas & Garcia Barriga. 9721 (COL, K, NY, 
P); Paramo de Guantiva, Alto de Canutos, 3200-3400 m, 3.x. 1940 (fl & fr), 
Cuatrecasas 10329 (COL, F, US). Paramo de La Rusia, NW.-N. de Duitama, 
Aislada, 3490 m, 8.xii.l972 (fl & fr), Cleef 6835 (U). Santander: Paramo de 
Santurban, near Vetas, 3950-4160 m, 17.1.1927 (fl), Killip & Smith 17567 (A, BM, 
K,US). 

Subsp. compactum differs from subsp. striatum in the subacute to acute, cucuUate, 
usually glandular leaves and the markedly ribbed sepals, both leaves and sepals being 
frequently red-tipped or red-tinged when young. It comprises two forms, which are 
geographically almost distinct and differ mainly in habit. The form in Santander 
('var. compactum'), which is confined to the Paramo de Santurban, has relatively 
stout stems with crowded branches that at higher altitudes (3570-4160 m) form a 
compact, caespitose clump or prostrate mat. The leaves are relatively stout, often 
broader towards the tip, and remain erect and distinctly tetrastichous, and the sepals 
are rarely red-tipped. Most of the Boyaca and Cundinamarca plants have taller, 
relatively slender, more loosely branched stems forming lax bushes ('var. gracile'), 
and the sepals are usually red with a hyaline margin. 

The 'gracile' form has been confused with the superficially similar Peruvian - 
Ecuadorian H. aciculare, from which it differs by its oblong (not lanceolate) sepals, 
which (like the leaves) are nearly always tipped or tinged reddish, become markedly 
ribbed and have relatively broad hyaline margins. In addition, the styles are capitate 
and the seeds only about half as long, and the leaves become outcurved, not twisted. 



49. Hypericum jaramilloi N. RoBson, sp. nov. 

H. caracasano Willd. affinis, sed foliis et floribus maioribus, sepalis longioribus, 
petalis latioribus, staminibus numerosioribus, stylis longioribus, stigmatibus angus- 
tioribus, differt. Type: Colombia, linea divisoria entre los Deptos. Santander del 
Norte y Cesar, entre Abrego y Las Jurisdicciones (Cerro de Oroque), 3440-3750 m, 
19-21. V. 1969 (fl), Garcia-Barriga & Jaramillo Meija 19810 (COL!, holotype). 

Shrub 1-1-5 m tall, erect, with branches strict, pseudo-dichotomous or occasionally 
lateral. Stems orange-brown, 4-angled when young, the subfoliar ridges broad, 
eventually terete, cortex exfoliating in strips; internodes 3-9 mm long. Leaves with 
± distinct petiole 1-3 mm long, ± densely imbricate at first, soon outcurving then 
usually widely spreading above petiole, subtetrastichous, deciduous above petiole 
without fading; lamina 9-15)-20) x 2-5-5(-6-5) mm, usually narrowly elliptic, 
plane to subincurved, not cucullate, midrib not prominent beneath, margin not 
distinct, concolorous, glaucous, coriaceous; apex acute, base angustate, incurved 
but not sheathing, pairs united to form narrow interfoliar ridge; basal vein 1, with 
3-4 pairs of ascending main laterals, obscurely branching distally, tertiary reticula- 
tion absent; laminar glands dense, sometimes obscure. Inflorescence 1 -flowered, 
with pseudo-dichotomous branches from node below; pedicel 5-13 mm long, slightly 
incrassate distally; upper leaves broader but not transitional. Flowers 25-30 mm in 
diam. , stellate. Sepals 9-14 x 3-6 mm, broadly to narrowly obovate or oblanceolate 
to elliptic-oblong or narrowly obovate or oblanceolate to elliptic-oblong or narrowly 
elliptic, acute to subacuminate, veins 5-7, unbranched, with midrib not or slightly 
prominent; glands linear to punctiform. Petals bright yellow, 12-17 x 6-10 mm, 
1-2-1-5 X sepals, obovate; apiculus acute; glands linear, distally punctiform. 
Stamens c. 70-80, longest 6-8 mm, c. 0-5 x petals. Ovary 3-5-4 x 2 mm, 
cylindric-ellipsoid; styles 3, 6-7 mm long, c. 1-5 x ovary, distally outcurved; stigmas 
scarcely capitate. Capsule c. 8 x 4-5 mm, cylindric-ellipsoid to cyHndric, shorter 
than sepals. Seeds not seen. 



THE GENUS HYPERICUM L. 

Paramo; 2600-3960 m. 

Costa Rica (Limon), Colombia (Magdalena, Cesar/Norte de Santander). Map 20. 

COSTA RICA. Limon: Cordillera de Talamanca, Cerro Kamuk massif, between 
Cerro Dudu and Cerro Apri, 2900-3100 m, 23 & 26.iii. 1984 (fl), Davidse, Herrera & 
Warner 25880 (MO). 

COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, transecto del Alto Rio 
Buritaca, 2900 m, 5.viii.l977 {^),Jaramillo etal. 5371 (BM, COL); loc. cit., 3300 m, 
9.viii.l977 (fl), Jaramillo etal. 5454 (COL, U). Cesar/Norte de Santander: Cerro de 
Oroque, Las Jurisdicciones, 3700-3900 m, 22-27.vii.1974 (fl), Garcia Barriga & 
Jaramillo 20598 (COL); Cerro de Oroque, 20 km al S. de Abrego, 3700-3960 m, 
19-21. vii. 1974 (fl), Garcia Barriga & Jaramillo 19728 (COL). 

H. jaramilloi is related on the one hand to H. pimeleoides and on the other to both H. 
cardonae and H. ruscoides. Its area of distribution overlaps that of H. pimeleoides in 
the Cerro de Oroque, where it can be distinguished by (i) the narrower spreading 
leaves with narrow (and usually longer) petioles and pinnate venation and (ii) the 
narrower sepals. From both H. cardonae and H. ruscoides it can be distinguished by 
size of parts (it resembles a larger, more primitive version of the latter), in particular 
by the longer styles. 



87 



50. Hypericum cardonae Cuatrec. 

in Revta Acad, colomb. Cienc. exact, fis. nat. 6: 62 (1944), in Trab. Com. Bot. etc. 
Colomb. : 36 (1944). Type: Venezuela, Tachira, Paramo de Tama, 3100-3300 m, 
vii. 1939 (fl), Cardona 303 (VEN!, holotype; F!, isotype). 

H. caracasanum sensu Turcz. in Bull. Soc. Nat. Moscou 31 (1): 389 (1858); Knuth in 
Reprium Spec. nov. Regni veg. Beih. 43: 483 (1927) pro parte, quoad spec, merid. , 
et auct. plur. 

H. caracasanum subsp. cardonae (Cuatrec.) N. Robson in Cleef, Veg. Pdram. 
Colomb. Cord. Or. : 305 (1981), comb, illegit. sine basionym. 

Shrub or shrublet 0-2-1 m tall, erect or rarely decumbent and rooting at the base, 
with branches strict, pseudo-dichotomous or lateral. Stems yellow- to orange-brown, 
4-lined and ancipitous when young, the subfoliar ridges broad, soon terete, cortex 
exfoliating in strips; internodes 3-7 mm long. Leaves with petiole 1-2-5 mm long. 




Map 20 49. H. jaramilloi A ; 50. H. cardonae •; 51. //. caracasanum: a. subsp. caracasanum ■, b. subsp. 
turumiquirense D . 





NORMAN K. B. ROBSON 

persistently imbricate-tetrastichous to widely spreading from above petiole, de- 
ciduous above petiole without fading; lamina 7-15 x 2-5-7 mm, broadly to narrowly 
elliptic or narrowly oblong, plane to incurved or subconduplicate, not cucullate, 
midrib not or slightly prominent beneath, margin plane to incrassate, concolorous, 
often glaucous to blue-green, coriaceous; apex acute to subrounded, base angustate 
to cuneate, the petioles not sheathing but united to form narrow interfoliar ridge; 
basal vein 1, with (l-)2-3 pairs of ascending lateral branches (often obscure), rarely 
visibly branching, not impressed beneath, tertiary reticulation not visible; laminar 
glands dense, visible beneath or not. Inflorescence 1-flowered, sometimes with 
several flowering branches clustered, with pseudo-dichotomous branches from one 
or two nodes below; pedicel 4-6 mm long, not incrassate distally; upper leaves not 
transitional. Flowers 12-18 mm in diam. , stellate or obconic. Sepals 6-12 x 2-4 mm, 
broadly to narrowly elliptic or oblong-elliptic to oblong-spathulate, acute to sub- 
acute ; veins 5-7 , sometimes branched distally , with midrib not or slightly prominent ; 
glands mostly linear, distally punctiform. Petals bright yellow to orange-yellow, 8-11 
X 3-6 mm, c. 1-3 x sepals, oblanceolate to obovate; apiculus acute; glands mostly 
linear, distally punctiform. Stamens 40-60, longest 4-7 mm long, c. 0-5 x petals. 
Ovary 2-5-3 x 1-5-2 mm, ovoid-ellipsoid to ellipsoid-subglobose; styles 3, 3-5 mm 
long, 1-1-6 X ovary, divergent to suberect; stigmas broadly capitate. Capsule 4-8 x 
2-4 mm, cylindric-ellipsoid, equalling or shorter than sepals. Seeds 1-1-2 mm long, 
ecarinate; testa finely scalariform. 

In humid (shaded or open) places in the paramo; 2500-4160 m. 

Costa Rica (Cartago, San Jose), Venezuela (Trujillo, Merida, Tachira, Apure), 
Colombia (Norte de Santander, Santander, Boyaca, Arauca). Map 20 (p. 87). 

COSTA RICA. Cartago: Cordillera de Talamanca, 'Cerro de la Muerte', Pan- 
American Highway 5 km above Millsville (8 km above Nivel), 3400-3300 m, vii.1949 
(fl). Holm & litis 454 (F, GH, K, MO, NY, P, U). San Jose: Valle de los Conejos 
(upper Rio Talari) and trails to Cerro Cirripo and Valle de los Lagos, 3400-3820 m, 
22.viii.1971 (fl), fiwrger 8281 (BM, DUKE, F). 

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, 3350 m, 
13.iii.l973 (fl), Cleef 91S2D (COL). Boyaca: paramos al NW. de Belen, cabeceras 
Quebrada Minas, Hoya Laguna El Alcohol, 3800 m, 26.ii.1972 (fl), Cleef 1S70 (U); 
Cocuy Mts, valley of Quebrada Playitas above Guican, 3800 m, 24. vi. 1984 (fl). 
Wood 4453 (K). Norte de Santander: Paramo de Tama, alredores de la Cueva, 3000- 
3200 m, 28.X.1941 (fl), Cuatrecasas, Schultes & Smith 12691 (COL). Santander: 
Paramo de Santurban near Vetas, 3950-4160 m, 17. i. 1927 (fl), Killip & Smith 
21167 (NY). 

VENEZUELA. Apure: Paramo de Pata de Judio, 30 km al S. de San Vicente de 
Revancha, 3000-3200 m, 19.1.1968 (fl), Steyermark & Dunsterville 101140 (NY, 
VEN). Merida: Paramo de Los Leones (La Lagunita, La Canada Grande), W. of 
Mucuruba, 3500 m, 31.V.1930 (fl & fr), Gehringer 138 (F, GH, NY, VEN); ViUa de 
Caracas, Las Lagunetas, xi. (fl & fr), Moritz 1167 (BM, GH, K, KW, S). Tachira: 
Paramo El Batallon, 3200-3600 m, 3.x. 1956 (fl), Vareschi 5491 (VEN); Hoya del Rio 
Tachira, cabeceras de la Quebrada de El Reposo, 2800-3000 m, 16.1.1973 (fr), 
Cuatrecasas, Ruiz-Terdn & Lopez-Figueras 28330 (BM, US); Paramo de Tama, 
cerca la frontera Colombo- Venezolana, 3000-3200 m, 20-23. v. 1967 (fl), Steyermark 
& Dunsterville 98735 (VEN). 

Some forms of H. cardonae from Merida are rather similar to H. caracasanum, but 
the capitate stigma and the thicker, glaucous leaves with glands obscure or invisible 
will easily distinguish it. There is a southward reduction trend in habit and in leaf 
shape, plants from the Paramo de Tama to Belen being dwarfer (almost suffruticose) 
and having smaller leaves, with thickened margin, obtuse to subrounded apex, 
and a longer thinner petiole. The Costa Rican population is more similar to 
plants from Merida, but shows a similar reduction tendency to that of the southern 
plants. 

For differences between H. cardonae and H. caracasanum, see the description of 
the latter species. The reduced form of//, pimeleoides can be distinguished from that 
of //. cardonae by the broader leaves with flabellate venation visible beneath. 



THE GENUS HYPERICUM L. 

5 1 . Hypericum caracasanum Willd . 

Sp. PL 3: 1458 (1802); Choisy, Prodr. monogr. fam. Hyperic. 58 (1821); Kunth in 
Humboldt, Bonpland & Kunth, Nova Gen. et Spec. PI. 5: 186 (1822); Choisy in 
DC. , Prodr. syst. nat. regni veg. 1: 554 (1824); Turcz. in Bull. Soc. Nat. Moscou 31 
(1): 389 (1858); R. Keller in Bull. Herb. Boissier II, 8: 177 (1908); in Engler & 
Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Knuth in Reprium Spec. nov. 
Regni veg. Beih. 43: 483 (1927) pro parte, excl. spec, merid.; Gleason in Bull. 
Torrey bot. Club 56: 103. (1929); Steyerm. & Huber, Fl. Avila: 493, t. 152B 
(1978). Type: Venezuela, Distrito Federal, Silla de Caracas, i.l800 (fl), Humboldt 
& Bonpland 649 (B, holotype, F!, US!, photographs; P!, isotype). 

Shrub orshrublet 0-5-1-8 m tall, erect, with branches strict, pseudo-dichotomous or 
lateral. Stems yellow- to orange-brown, 4-6-lined and compressed when young, the 
subfoliar ridges broad, soon or eventually terete, cortex exfoliating in strips; 
internodes 1-4 mm long. Leaves subsessile to broadly or narrowly petiolate with 
petiole up to 1-5 mm long, ± densely imbricate, tetrastichous, ascending to outcurv- 
ing, deciduous above petiole without fading; lamina 5-14(-17) x 2-7 mm, ± 
narrowly elliptic or ovate-elliptic to oblong, plane to incurved or slightly cucuUate, 
midrib slightly prominent to impressed beneath, margin plane to recurved or slightly 
incrassate, paler beneath or concolorous or with margin pinkish, not glaucous, 
coriaceous; apex acute to subacute, base angustate, the petioles not or scarcely 
sheathing but united to form narrow interfoliar ridge; basal vein 1, with 1-3(4) pairs 
of ascending lateral branches, not visibly branching, not impressed beneath, tertiary 
reticulation not visible; laminar glands dense, usually visible beneath. Inflorescence 
1-flowered, with pseudo-dichotomous branches from node below; pedicel 2-5-5-5 
mm long, not or scarcely incrassate upwards; upper leaves not transitional. Flowers 
12-20 mm in diam., stellate. Sepals 5-l\ x 2-4-5 mm, broadly or narrowly elliptic or 
oblong-elliptic to obovate-spathulate or oblanceolate, acute to subacute; veins 3-7, 
sometimes branched distally, with midrib slightly prominent; glands linear, distally 
punctiform. Petals bright? to deep yellow, 8-13 x 3-5-6 mm, 1-4-2 x petals, 
oblanceolate; apiculus acute; glands mostly linear, distally interrupted to punct- 
iform. Stamens c. 50-70, longest c. 4-7 mm long, c. 0-5 x petals. Ovary 2-3-5 x 2 
mm, ovoid-ellipsoid; styles 3, 4-5 mm long, c. 1-5-2-5 x ovary, diverging and 
sometimes incurved; stigmas narrow or clavate. Capsule (5)6-8 x (2)3-4 mm, 
cyhndric-ellipsoid, equaUing sepals. Seeds c. 1-2 mm long, ecarinate; testa finely 
scalariform. 

In humid or shaded habitats in the paramo and subparamo; (1600) 2000-2765 m. 

Venezuela (Distrito Federal, Miranda, Aragua, Anzoategui, Sucre). Map 20 (p. 87). 

H. caracasanum and H. cardonae are vicarious species which differ in leaf shape, 
thickness and glandularity, and in stigma shape, but H. caracasanum is more 
restricted in morphological variation and distribution than is its vicariad. In H. 
caracasanum the leaf glands are always visible and the stigma is not or scarcely 
enlarged. It appears to be directly related to H. jaramilloi. 

The extreme easternmost population of//, caracasanum, in the Cerro de Turumi- 
quire (Sucre), has smaller, more crowded leaves and somewhat smaller flowers than 
the rest of the species, thus showing a trend in morphology as well as geography 
towards //. ekmanii from Hispaniola. The variation between the Sucre population 
and the rest is apparently almost continuous, and therefore subspecies is the 
appropriate rank for the two populations. 

51a. H. caracasanum subsp. caracasanum 

Brathys caracasana (Willd.) Spach, Hist. nat. veg. Phan. 5: 448 (1836) in Annls Sci. 
nar. (Bot.) II, 5: 366 (1836). 

Icones: Steyerm. & Huber, Fl. Avila: t. 152B (1978); Fig. 13A. 

Shrub or shrublet 0-5-1 m tall. Stem internodes 2-4 mm long. Leaves subsessile or 
with petiole up to 1-5 mm long; lamina 7-14(-17) x (2-)3-5-7 mm, narrowly elliptic 



89 





90 






NORMAN K. B. ROBSON 

to oblong. Pedicels 4-5-5-5 mm long. Flowers 15-20 mm in diam. Sepals 8-11 x 
2-4-5 mm, broadly or narrowly elliptic or oblong-elliptic to obovate-spathulate, 
veins 5-7. Petals bright? yellow, 11-13 x 5-6 mm, c. 1-4 x sepals. Ovary 2-3 x 
1-5-2 mm; styles 4-4-5 mm long, c. 1-5-2 x ovary. 

In humid habitats in the paramo; (1600)2000-2763 m. 

Venezuela (Distrito Federal, Miranda, Aragua, Anzoategui). 

VENEZUELA. Anzoategui: Rio Mantiales E. of Bergantia, Cerro Peonia above 
Santa Cruz, 2350 m, 21.iii.l945 (fl), Steyermark 61667 (F, NY, S, VEN). Aragua: 
prope Colonia Tovar, 1854-1855 (fl), Fendler 40 (GH, K, MO, P). Distrito Federal: 
Silla de Caracas, Cordillera de Avila, between Los Venados and Pico Oriental, 
1675-2640 m, 27-28. ii. 1944 (st), Steyermark 55644 (F, NY). Miranda: Pico de 
Naiguata, above Los Chorros, 2200-2675 m, 16-17. vi. 1945 (fl), Steyermark 62943 
(F, NY). 

51b. H. caracasanum subsp. turumiquirense (Steyerm.) N. Robson, stat. 
nov. 

H. caracasanum var. turumiquirense Steyerm. in Fieldiana Bot. 28: 393 (1952). Type: 
Venezuela, Sucre, Cerro Turumiquire, north-facing slopes, 2360-2500 m, 
6.V.1945 (fl), Steyermark 62579 (F!, holotype; NY!, US!, isotypes). 

Shrub 0-8-1-8 m tall. Stem internodes 1-2 mm long. Leaves with petiole 1-1-5 mm 
long; lamina 5-8 x 2-3-5 mm, ovate-elliptic to elliptic. Pedicels 2-5-3 mm long. 
Flowers 12-18 mm in diam. Sepals 5-6 x 2-3 mm, obovate-oblong to oblanceolate- 
spathulate, veins 3-5. Petals deep yellow, 8-12 x 3-5-4-5 mm, 1-5-2 x sepals. 
Ovary c. 2 x 1 mm; styles 4-5 mm long, 2-2-5 x ovary. 

On steep north-facing sandstone slopes in subparamo vegetation; 2360-2910 m. 

Venezuela (Sucre). Confined to the Cerro Turumiquire. 
VENEZUELA. Sucre: Cerro de Turumiquire, 1925 (fl), Tate 329 (US). 

52. Hypericum ekmanii A. H. Liogier 

in Phytologia 47: 183 (1980). Type: Dominican Repubhc, Peravia, San Jose de Ocoa, 
2000 m, 4.vi.l978 (fl & iT),A. & P. Liogier 21672 (SDM, holotype; GH!, isotype). 

Icon: Fig. 13B. 

Shrub up to 0-75 m tall, erect, forming thickets, with branches ascending, pseudo- 
dichotomous and sometimes lateral. Stems orange-brown, 4-lined and compressed 
when young, the subfoliar ridges broad, soon 2-lined, eventually terete, cortex 
exfoliating at first in strips between subfoliar ridges; internodes 2-4 mm long. Leaves 
with short petiole (0-5-1 mm long), densely spreading or imbricate, tetrastichous, 
deciduous above petiole without fading; lamina 6-10 x 2-4 mm, eUiptic, plane to 
slightly concave, scarcely cucullate, midrib not or sHghtly prominent beneath, 
margin incurved, not incrassate, concolorous, not glaucous or slightly so beneath, 
subcoriaceous; apex acuminate to apiculate, base angustate to cuneate, the petioles 
not sheathing but united to form very narrow interfoliar ridge; basal vein 1, with 1 
short near-basal pair of branches or unbranched; laminar glands dense, visible on 
both sides. Inflorescence 1 -flowered, with pseudo-dichotomous branches from node 
below; pedicel 3-10 mm long, not incrassate upwards; upper leaves not transitional. 
Flowers c. 12-15 mm in diam., stellate. Sepals 5-6 x 1-2-5 mm, unequal to equal 
narrowly oblong to oblanceolate-spathulate, acute to subacuminate; veins 3, un- 
branched, with midrib not prominent, glands linear, punctiform in distal 1/4. Petals 
deep? yellow, 8-10 x 2-4 mm, c. 1-5 x sepals, oblong-oblanceolate; apiculus 
subacute; glands nearly all punctiform. Stamens c. 80, longest 5-6 mm long, c. 0-6 x 
petals. Ovary 2x1 mm, narrowly ovoid; styles 3, 5-6 mm long, 2-5-3 x ovary, 
diverging-incurved; stigmas clavate. Capsule 3-6 x 2-3 mm, ellipsoid, shorter than 
sepals. Seeds 1-2 mm long, ecarinate; testa finely ribbed-scalariform. 

On rocky slopes in cloud forest; 1300-2550 m. 



THE GENUS HYPERICUM L. 



91 




Fig. 13 A. H. caracasanum subsp. caracasanum: (a) habit; (b) leaf; (c)- sepal; (d) petal; (e) stamens 
(partly cut away) and ovary. B. H. ekmanii: (f) habit; (g) stamens (partly cut away) and ovary. C. H. 
pycnophyllum: (h) habit. D. H. millefolium: (i) habit (a, f, h, i x Vi; b x 2; c-e, g x 3). A. Davidse & 
Gonzalez 19550; B. Ekman 11718; C. Liogier 21790; D. Holdridge 1776. 



Dominican Republic (Sierra de Ocoa). Map 21 (p. 92). 

DOMINICAN REPUBLIC. Azua: Sierra de Ocoa, San Jose de Ocoa, S. slope of 
Tetero de Majia, c. 2550 m, l.ii.l929 (fl & fr), Ekman H. 11718 (S, US). 

Ekman (on the back of the label) states that the species is quite common in 'these 
mountains' (i.e. Sierra de Ocoa) and occurs at 1300 m near Bejucal. The gap in 
distribution between H. ekmanii (Hispaniola) and its nearest relative, H. caracasa- 
num subsp. turumiquirense (NE. Venezuela), is likely to have resulted from long- 
distance dispersal and therefore cannot be adduced to support the theory that North 
and South America were originally linked via the Greater Antilles and the Yucatan 
Peninsula (see p. 9). 



92 



NORMAN K. B. ROBSON 





\J 






v*^ 




-^r 


'^^^^^T^ 




r' (^ 




1 1 ^ 



Map 21 52. //. ekmanii • ; 53. //. pycnophyllum A ; 54. //. millefolium 



-v Q--: 



5xi5 



53. Hypericum pycnophyllum Urban 

Symb. Antill. 7: 523 (1913). Type: Dominican Republic, La Vega, Loma Rosilla, 
1700 m, 6.vii.l912 (fl), Fuertes 1749 (Bf, holotype; NY!, P!, W!, isotypes). 

Icon: Fig. 13C. 

Shrub 0-2-1 m tall, erect, with branches very strict, pseudo-dichotomous and lateral. 
Stems reddish-brown, 2-lined and compressed when young, the subfoliar ridges 
broad, very soon terete, appearing articulated when old, cortex exfoliating at first in 
strips between subfoliar ridges; internodes 1-1-5 mm long. Leaves petiolate with 
short petiole (c. 0-5 mm long), densely imbricate, markedly tetrastichous, deciduous 
above petiole without fading or with attached cortex below after fading; lamina 3-9 
X 0-5-2 mm, very narrowly elliptic to linear, incurved to canaliculate, ± cucuUate, 
midrib impressed beneath, margin narrowly hyaline, dull above, lucent beneath, not 
glaucous, coriaceous; apex acute, base angustate to above 'articulation' then 
broadening, pairs forming narrow interfoliar ridge; basal vein 1, unbranched; 
laminar glands dense, obscure, visible above only. Inflorescence 1-flowered, terminal 
and on lateral branches, sometimes with single or pseudo-dichotomous branches 
from node below; pedicel absent or up to 3 mm long, not or slightly incrassate 
upwards; upper leaves not transitional. Flowers 10-20 mm in diam., stellate to 
obconic. Sepals 5-6(-7) x 1-2 mm, subequal, lanceolate to oblong-lanceolate, 
acute to acuminate, cucullate, with margin narrowly hyaline; veins 3-7, unbranched, 
midrib not prominent; glands Hnear, uninterrupted. Petals deep? yellow to orange- 
yellow, 7-14 X 3-5-7 mm, c. 1-6 x sepals, oblanceolate; apiculus acute; glands 
linear, distally interrupted. Stamens 55, longest 4-5-6-5 mm long, c. 0-6 x petals. 
Ovary c. 2 x 1 mm, ovoid; styles 3, 3-5-4 mm long, 1-5-2 x ovary, diverging- 
incurved; stigmas subclavate. Capsule 5-7 x 2-5-3 mm, narrowly ovoid, shorter 
than sepals. Seeds c. 1 mm long, ecarinate; testa finely ribbed-scalariform. 

In open Pinus occidentalis forest and among rocks at higher altitudes; 1700-3175 m. 

Dominican Repubhc (Santiago, La Vega, San Juan, Azua). Map 21. 

DOMINICAN REPUBLIC. Azua: El Pico del Yaque, 3125 m, 3. i. 1944 (st), 
Jimenez 232 (US). La Vega: Sabana Alte, c. 2700 m, 16.X.1929 (fl & fr), Ekman H. 
13793 (S). San Juan: Pico Duarte, 3170 m, 10-14.vii.l976(fl), A. c& P. Liogier 25040 
(NY). Santiago: Moncion, top of Cerro Lucio, c. 2525 m, 12. vi. 1929 (fl & fr), Ekman 
H.12850 (C, F, K, NY, S, US). 

The specimens from higher altitudes have shorter leaves. Hypericum pycnophyllum 
is apparently more widespread than H. ekmanii and links it morphologically with H. 
millefolium. 



THE GENUS HYPERICUM L. 



93 



54. Hypericum millefolium Urban & Ekman 

in Ark. Bot. 20 A, no. 15: 79 (1926). Type: Haiti, Massif de Selle, Morne de la Selle, 
Morne Emerillon, 2000 m, i.l925 (fl & fr), Ekman H. 3126 (Bt, holotype; K!, 
NY!,S!,isotypes). 

Icon: Fig. 13D. 

Shrub up to c. 2 m tall, erect, juniperoid, with branches strict, pseudo-dichotomous 
(extension) and shorter, lateral (flowering). Stems 4-6-lined and ancipitous when 
young, persistently 4-angled and ancipitous until cortex exfoliates at first in strips 
between subfoliar ridges, then terete, fragile, green becoming pale grey; internodes 
1-2 mm long. Leaves sessile, imbricate to curved-ascending, tetrastichous, eventu- 
ally deciduous with attached cortex below after (?) fading; lamina 0-7-2 x 0-8-1 
mm, lanceolate-triangular, incurved, markedly cucuUate, midrib impressed be- 
neath, margin narrowly hyaline, dull above, lucent or ± glaucous beneath, 
coriaceous; apex acute, pungent, base broad, subamplexicaul, pairs united to form 
narrow interfoliar ridge; basal vein 1 , unbranched; laminar glands obscure. Infloresc- 
ence 1-flowered, terminal and on numerous short lateral branches (branching or 
not), occasionally with pseudo-dichotomous branches from node below; pedicel 
absent or up to 0-5 mm long, incrassate upwards; upper leaves not transitional. 
Flowers 5-7 mm in diam., obconic. Sepals 3-3-5 x 1 mm, lanceolate, acute and 
incurved-cucullate to pungent; veins 5, unbranched, midrib ± prominent; glands 
Hnear, uninterrupted. Petals deep? yellow, 6-7 x 2-5-3 mm, 2 x sepals, oblan- 
ceolate; apiculus very acute; glands linear, distally subpunctiform. Stamens c. 50, 
longest 4-5 mm long, 0-6-0-7 x petals. Ovary 1-5x1 mm, broadly ovoid; styles 3, 
4-5 mm long, c. 3 x ovary, diverging-incurved; stigmas narrow. Capsule 4x2 mm, 
ovoid, exceeding sepals. Seeds not seen. 

Among steep limestone rocks; 1750-2350 m. 

Haiti (Massif de la Selle). Map 21. 

HAITI. Quest: Massif de la Selle, Morne la Selle, Bouche Bouqui, 1750 m, 
15.X.1943 (fl & fr), Holdridge 1776 (BM, MICH, NY, US); Croix des Bouquets, 
Morne Badeau, c. 2200 m, 24.ii.1927 (fl), Ekman H. 7679 (F, S, US), Petionville, top 
of Morne Cabaio, 2350 m, 10.iv.l927 (fl), Ekman H. 7979 (C, S). 

Hypericum millefolium, with its cricoid habit, is at first glance quite distinct from any 
other species in sect. Brathys. Closer investigation, however, shows it to be similar to 
the smaller form of H. pycnophyllum. Its triangular leaves and numerous fragile 
flowering stems, nevertheless, make it easily recognizable. 



(fi 




55. Hypericum ruscoides Cuatrec. 

in Ciencia Mex. 4: 63 (1943). Type: Colombia, Putumayo, Paramo de San Antonio 
del Bordoncillo, between El Encano and Sibundoy, 3250 m, 4. i. 1941 (fl), Cuatre- 
casas 11720 (COL, holotype; F!, P!, US!, isotypes). 

Shrub 0-1-1-5 m taU, erect or with stems sometimes decumbent and rooting at the 
base, with branches strict, pseudo-dichotomous and sometimes lateral. Stems yel- 
low-brown, 4-lined and compressed when young, the subfoliar ridges broad, even- 
tually terete, cortex exfoliating in strips between subfoliar ridges; internodes 2-5-6 
mm long. Leaves petiolate with petiole 1-5-3-5 mm long, ascending to spreading, 
tetrastichous, deciduous above petiole without fading; lamina 8-18 x 3-6(-8) mm, 
narrowly ovate or lanceolate to narrowly oblong-elliptic or the uppermost elliptic, 
plane or slightly conduplicate, not or rarely carinate, slightly cucuUate, midrib not or 
slightly impressed beneath, margin not distinct or rarely subincrassate, concolorous, 
sometimes ferrugineous, ± lucent or dull and undulate-papillose on both sides, 
sometimes glaucous, coriaceous; apex acute to apiculate, base angustate, the 
petioles not sheathing but united to form narrow interfoliar ridge; basal vein 1, with 
2-4 pairs of obscure ascending branches, rarely visibly branched or reticulating; 
laminar glands dense, sometimes visible on either or both sides. Inflorescence 
1-flowered, with pseudo-dichotomous branches from node below; pedicel 4-8 mm 




94 




NORMAN K. B. ROBSON 

long; upper leaves often broader but not transitional. Flowers 15-25 mm in diam., 
stellate. Sepals 5-10 x 1-5-4 mm, obovate-spathulate to narrowly oblong (unequal 
to subequal), acute to obtuse; veins 5, branched or not, midrib not prominent; glands 
linear, distally punctiform. Petals deep yellow to orange-yellow, 8-15 x 4-6 mm, c. 
1-5 X sepals, obovate-spathulate; apiculus subacute to obtuse; glands linear, distally 
punctiform. Stamens 30-60, longest 4-6-5 long, c. 0-5 x petals. Ovary 2-3 x 1-5-2 
mm, ovoid; styles 3, 2-5-4 mm long, c. 1-3 x ovary, suberect to divergent; stigmas 
broadly capitate. Capsule 4-8 x 3-3-5 mm, cylindric-ellipsoid to ellipsoid, shorter 
than sepals. Seeds c. 1-5 mm long, ecarinate; testa finely scalariform. 

In paramo; 2450-3800 m. 

Colombia (Cordilleras Oriental, Central and Occidental from Boyaca and Cundina- 
marca to Putumayo), Ecuador (Pichincha and Napo: Volcan Cayambe). Map 22. 

COLOMBIA. Boyaca: Villa de Leiva, Parque Igaque, Quebrada Carrizal, 3400 m, 
25. xi. 1984 (fl & fr). Wood 4621 (K). Caldas: Navarco, 3000 m, 12.ix.l951 (fl), 
Romero-Castaneda 3021 (COL). Cauca: Cordillera Central, Paramo de las Moras, 
3800-3700 m, 19.ii.l969 (fl), Cuatrecasas 27424 (BM, COL, US). Cundinamarca: 
Municipio de Fomeque, paramo de Chingaza, cerros vecinos a la Laguna y limites 
con el Depto. del Meta, 3300 m, 2.xi.l966 (fl), Huertas & Camargo 6572 (COL). 
Huila: Cordillera Oriental, 20 km SE. of Gigante, 20.ix.l944, Little 8687 (COL). 
Meta: Paramo de Sumapaz, Cerro Nevado de Sumapaz, 3725 m, 29. i. 1972 (fl), Cleef 
1206 (BM, COL, U). Nariiio: Pasto a Sibundoy, c. 3000 m, 5. i. 1952 (fl), Fernandez & 
Knoth 1023 (COL). Putumayo: Valle de Sibundoy, Sibundoy, 2800 m, 30.vi.l953 
(fl), Schultes & Cabrera 20138 (GH, NY). Tolima: Cordillera Central, Municipio de 




Map 22 55. H. ruscoides A ; 60, H. selaginella O. 



THE GENUS HYPERICUM L. 

Roncevalles, paramo de Normandia, 3200 m, 8.xi.l980 (fl), Camargo 7508 (COL). 
Valle: Cordillera Occidental, Los Farallones, cerro Alto de Buey, 3500-3600 m, 
n.x.l944, Cuatrecasas 17917 (BM, F, US), 17940 ,;COL). 

ECUADOR. Pichincha: Cordillera Oriental, Laguna de Chiqui, NE. of 
Cayambe, 3800 m, 4.vi.l944 (fl), Acosta Solis 8151 (F). Napo: ENE. of Cayambe 
Mtn, 3210 m, 6.xii.l961, Cazalet & Pennington 5522 (K, NY). 

Hypericum ruscoides is related to H. jaramilloi, being intermediate between it and 
H. lancioides. It can easily be confused with reduced forms of H. pimeleoides in the 
area where the ranges of these species overlap, but can be recognized by the usually 
narrower and less crowded leaves with longer, narrower petioles, acute apex, and no 
free veins, i.e. the venation is pinnate. 

H. ruscoides appears to be rare in the Cordillera Occidental. Two collections from 
Valle de Cauca, Los Fardallones, cerro Alto del Buey {Cuatrecasas 17917, 17940) 
have thinner, narrower, more shortly petiolate leaves than usual, and are nearest in 
form to H. lancioides. Cuatrecasas has distinguished them as 'var. occidentale' on the 
labels, and if these characters are constant in Cordillera Occidental populations, 
then this population may warrant recognition as a subspecies. 



95 



56. Hypericum llanganaticum N. Robson, sp. nov. 

H. struthiolifolio Juss. affinis, sed statura alta, ramis haud radicantibus, foliis planis 
vel incurvatis, manifeste tetrastichis, plusminusve glaucis, sepalis oblongo- 
oblanceolatis haud porcatis, differt. Type: Ecuador, Napo, Llanganati, paramo SE. 
of Chosa Aucacocha, between Aucacocha and Pan de Azucar, 3800-3900 m, 
15.V.1982 (fl), 0llgaard, Holm-Nielsen etal. 38464 (AAU!, holotype). 

Shrub 0-4-2 m tall, erect to decumbent but not rooting, with branches strict, 
pseudo-dichotomous or lateral. Stems orange-brown, 4-lined and ancipitous when 
young, eventually terete, cortex exfoliating in irregular flakes; internodes 1-2 mm 
long. Leaves sessile or rarely subsessile, erect to narrowly spreading, ± imbricate- 
tetrastichous, deciduous above the base after fading; lamina 7-13 x 1-5-4 mm, 
narrowly elliptic to linear-oblong, plane or slightly incurved, scarcely to markedly 
cucullate, midrib impressed beneath, margin not or very narrowly hyaline, concol- 
orous, rather glaucous, ± coriaceous; apex acute, base angustate, not sheathing, 
pairs united to form rather shallow interfoliar ridge; basal vein 1, (always?) unbran- 
ched; laminar glands dense, visible above and rarely also beneath; marginal glands 
rather dense. Inflorescence 1-flowered, with pseudo-dichotomous branches from 
node below; pedicel 3-4 mm long; upper leaves not transitional. Flowers c. 20-25 
mm in diam., stellate. Sepals 7-5-9 x 1-8-2-3 mm, oblong-oblanceolate, acute, not 
or slightly incurved-cucullate; veins 5, branched, not or scarcely prominent beneath; 
glands striiform to punctiform. Feffli/5 (bright?) yellow, 12-16 x 4-10 mm, c. 1-5-1-8 
X sepals, oblong-oblanceolate; apiculus acute; glands linear to striiform. Stamens c. 
80-90, longest 6-10 mm long, 0-5-0-7 x petals. Ovary c. 3 x 2-5 mm, broadly ovoid; 
styles 3, 3-6-4-3 mm long, 1-2-1-4 x ovary, divergent; stigmas not or narrowly 
capitate. Capsule c. 5-5 x 3-5 mm, ovoid-pyramidal, shorter than sepals. Seeds not 
seen. 

In grassy paramo with scrub or reed forest; 3250-4200 m. 

Ecuador (Tungurahua, Napo). Confined to the Cordillera de Los Llanganates. Map 
23 (p. 96). 

ECUADOR. Napo: Cordillera de Los Llanganati, Chihuila o Ainchilibi, 3750 m, 
25-29. viii. 1959 (fl), Barclay & Juajibioy 8996 (MO, NY); junction of Rio Chalupas 
and Rio Agua Buenas, E. end of upper Chalupas valley, 3250 m, 11. viii. 1979 (fl), 
Holm-Nielsen 19074 (AAU). Tungurahua: Cordillera de Llanganates, near Las 
Torres, 3700-3800 m, 22. xi. 1939 (fl), Asplund 9878 (S); Llanganati Mts., Lake 
Aucacocha, 3700 m, vii.1969 (st), Edwards 35 (K). 

H. llanganaticum comprises a relict population intermediate in form and distribution 
between H. ruscoides and H. struthiolifolium. Asplund 9878 differs from the other 
specimens in its broader leaves and thus approaches H. ruscoides, differing from it 




96 



NORMAN K. B. ROBSON 




Map 23 56. H. llanganaticum D; 57. //. struthiolifolium 



essentially only in the narrower leaves with a slightly cucullate apex and narrower 
stigmas. The remaining specimens are nearer to H. struthiolifolium but differ from it 
in having more crowded tetrastichous leaves (which are dull blue-green or glaucous, 
not lucent) and unribbed, oblanceolate to oblong sepals. 

57. Hypericum struthiolifolium Juss. 

in Annls Mus. Hist. nat. Paris 3: 161, t. 16, f.2 (1804) ['struthiolae folium']; Choisy, 
Prodr. monogr. fam. Hyperic. : 58 (1821), in DC. , Prodr. syst. nat. regni veg. 1: 553 
(1824); R. Keller in Bull. Herb. Boissier II, 8: 176 (1908), in Engler & Prantl, Nat. 
Pflanzenfam. 2nd ed. 21: 181 (1925). Type: Peru, without precise locahty or date 
(fl & fr), Dombey s.n. (P-JUS!, holotype; F!, P!, isotypes). 

Brathys struthiolifolia (Juss.) Spach, Hist. nat. veg. Phan. 5: 449 (1836) 
['struthiolaefolia'], in Annls Sci. nat. II, 5: 366 (1836). 

H. struthiolifolium var. genuinum Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18: 
293 (1862) ['struthiolaefolium'], pro parte excl. spec. cit. 

H. weberbaueri R. Keller in Bot. Jahrb. 42: 129 (1908), in Bull. Herb. Boissier, II, 8: 
178 (1908), in Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Gleason 
in Bull. Torrey bot. Club 56: 102, 103, 105 (1929), pro parte quoad typum. Type: 
Peru, Junin, Tarma, Huacapostana, 3000-3100 m, n.d. (fl & fr), Weberbauer 2b9?> 
(Bt, holotype; F!, GH!, NY!, US!, photographs; G!, isotype). 

Shrub up to c. 0-5 m tall, erect or sometimes decumbent and rooting, with branches 
strict, pseudo-dichotomous or sometimes lateral. Stems orange-brown, 4-lined and 
ancipitous when young, eventually terete, cortex exfoliating in irregular flakes; 
internodes 2-7 mm long. Leaves sessile, suberect to spreading or slightly outcurving, 
not or scarcely imbricate, deciduous above the base after fading; lamina 8-15 x 
1-5-2-5 mm, very narrowly elliptic to Hnear-oblong, ± incurved- cucullate, midrib 



THE GENUS HYPERICUM L. 

mostly impressed beneath, margin not or slightly hyahne, concolorous, lucent, not 
glaucous, ± coriaceous; apex acicular to subacute, base angulate, not sheathing, 
pairs united to form shallow interfoliar ridge; basal vein 1, occasionally with 1-2 
pairs of obscure branches; laminar glands rather dense, visible above but not 
beneath. Inflorescence 1-flowered, with pseudo-dichotomous branches from node 
below; pedicel 5-9 mm long; upper leaves not transitional. Flowers 15-25 mm in 
diam., stellate. Sepals 5-8 x 1-3 mm, ovate-lanceolate to oblong, acute, incurved- 
cucullate; veins 5-9, not branched, prominent beneath; glands linear, punctiform in 
distal third. Petals bright yellow, (8-)10-15 x 3-7 mm, c. 1-5 x sepals, oblong- 
oblanceolate to oblong-obovate; apiculus acute; glands striiform. Stamens 60-75, 
longest 5-9 mm long, 0-5-0-6 x petals. Ovary 2-5-3-5 x 1-5-2 mm, broadly ovoid; 
styles 3(4), 3-4 mm long, 1-3-1-5 x ovary, spreading; stigmas not or narrowly 
capitate. Capsule 4-5 x 2-2-5 mm, ellipsoid, shorter than sepals. Seeds c. 0-7 mm 
long, ecarinate; testa finely scalariform. 

On open hillsides among shrubs; 3000-3700 m. 

North and Central Peru (Amazonas, Pasco, Junin, Huancavelica, Cuzco). Map 23. 
PERU. Amazonas: Chachapoyas, Pass of Piscohuaiiuna, 2700 m, viii.1938 (fl), 
Sandeman 239 (K); Bagua, Cordillera Colan E. of La Peca, 2880 m, 8.viii.l978 (fl), 
Barbour 3230 (MO). Cuzco: Urubamba, Puyupatamarca area, 3200 m, 29.iii.1894 
(fl), Vargas 2762 (F). Huancavelica: Tayacaja, between Marcavalle and Huachocol- 
pa, 3600 m, 24.iv.1963 (fl & fr), Rovar 4225 (NY). Junin: Chihfruta to Huancayo, 11 
km, 3700 m, 30.x. 1968 (fl), B. & C. Maguire 61657 (BM, NY). Pasco: Oxapampa, 
Cordillera Yanachaga 12 km SE. of Oxapampa, 2700-2800, 7.X.1982 (fl), Foster 
9048 (BM, MO). 

H. struthiolifolium was one of the first species in sect. Brathys to be described, and 
the name has frequently been misapplied. The species is confined to north and 
central Peru, where it is apparently relatively uncommon. It differs from H. 
llanganaticum in the less crowded, more incurved, lucent (not glaucous) leaves and 
the usually smaller flowers with sepals that are ribbed and usually broader below the 
middle. Also, its decumbent stems sometimes root. Its distribution overlaps con- 
siderably that of the derivative H. andinum, but these species remain distinct. 



97 




58. Hypericum andinum Gleason 

in Torreya 29: 137 (1929). Type: Bolivia, Mt. Tunare, 1891 (fl & fr), Bang 1026 (NY!, 
holotype; BM!, F!, GH!, Kl, MO!, US!, W!, isotypes). 

H. struthiolifolium var. parviflorum R. Keller in Bull. Herb. Boissier II, 8: 182 (1908) 
['struthiolaefolium']. Type: Bolivia, La Paz, Larecaja, Majabaga, Catarguata, 
2800-3200 m, x.l858-v.l859 (fl & fr), Mandon790 (G!, holotype; BM!, GH!, K!, 
NY!, P!,S!,W!, isotypes). 

Shrub or shrublet 0-1-0-5 m tall, erect or decumbent and rooting, bushy with 
branches strict, pseudo-dichotomous or lateral. Stems reddish to yellowish-brown, 
4-angled when young, soon 2-lined then terete, cortex exfoliating in irregular flakes; 
internodes 1-4 mm long. Leaves sessile, suberect to outcurved, subimbricate to 
spreading, deciduous above the base after fading; lamina 3-5-10 x 0-8-1 -5 mm, 
narrowly oblong-elliptic to linear, incurved-cucullate, midrib mostly impressed 
beneath, margin ± narrowly hyaline, otherwise concolorous, not glaucous, sub- 
coriaceous to chartaceous; apex acute to rounded, base cuneate to parallel, not 
sheathing, pairs united to form shallow interfoliar ridge; basal vein 1, unbranched; 
laminar glands dense, impressed, visible beneath. Inflorescence 1-flowered, with 
pseudo-dichotomous branches from node below; pedicel 3-6 mm long; upper leaves 
not transitional. Flowers 10-18 mm in diam., stellate. Sepals 3-5-7 x 0-8-2 mm, 
broadly to narrowly lanceolate, subacuminate to acute, cucullate, with hyaline 
margin; veins (3)5(7), unbranched, ± prominent; glands linear, punctiform, in distal 
third. Petals golden yellow, 6-10 x 3-5 mm, c. 1-5 x sepals, narrow obovate to 
obovate-oblong; apiculus acute to obtuse or obsolete; glands shortly striiform to 
punctiform. Stamens 20-50, longest 3-6 mm long, c. 0-5 x petals. Ovary 1-8-2-3 





98 



NORMAN K. B. ROBSON 



11/ 



(-3-3) X 1-1-2 mm, narrowly ovoid; styles 3, 1-5-2-5 mm long, 0-6-0-7 x ovary, 
free, spreading; stigmas broadly capitate. Capsule 4-6 x 1-2-3 mm, ovoid, shorter 
than sepals. Seeds not seen. 

On open or stony grassy slopes and Sphagnum bogs, sometimes near the snowline; 
2400-4200 m. 

Peru (La Libertad, Junin, Huancavelica, Cuzco, Puiio), Bolivia (La Paz, Cochabam- 
ba, Santa Cruz). Map 24. 

PERU. Cuzco: Paucartambo Valley, 26 km from Cuzco to Pilcopata, 3425 m, 
27.ix. 1968 (fl), B. & C. Maguire 61564 (BM, NY). Huancavelica: Tayacaja, between 
Colcabamba and Paucarbamba, Arriba de la Hda. Jocas, 3900 m, 20. iv. 1954, Tovar 
2020 (NY). Junin: Huancayo, ii.l939 (fl), Soukup 389 (F). La Libertad: Huamachu- 
co, Jalca de Huaguil, 3950 m, 24.vi.1958 (fl), Lopez & Sagdstegui 2736 (NY). Puiio: 
Ayapata, vi.l854 (fl & fr), Lechler 1817 (K, P, W). 

BOLIVIA. Cochabamba: Road to Chapare [from Cochabamba], Km 104, 3100 
m, 23.xii.1926, Steinbach 650 (F, MO, NY, S, U, US, WIS). La Paz: Sorata, 3900 m, 
ii.l886 (fl & fr), Rusby 1350 (MICH, NY). Santa Cruz: Comarapa, Cerro de San 
Mateo, 3400 m, 25.x. 1928 (fl & fr), Steinbach 8535 (BM, F, GH, K, MO, NY, S, U). 

Although the area of//, andinum overlaps considerably that of//, struthiolifotium, 
the former can be distinguished by the shorter capitate styles and the narrower, 
usually less acute, recurving leaves with markedly hyahne margins. 

59. Hypericum lancioides Cuatrec. 

in Ciencia Mex. 4: 65 (1943). Type: Colombia, Putumayo, Paramo de San Antonio 
del Bordoncillo, between El Encano and Sibundoy, 3250 m, 4. i. 1941, Cuatrecasas 
11721 (COL, holotype; F!, US!, isotypes). 

Shrub c. 0-1-1 m tall, erect and bushy or ± decumbent and caespitose, branches 
strict, pseudo-dichotomous and lateral, or spreading (very rarely pinnate). Stems 
orange-brown, 6-lined and compressed when young, the subfoliar ridges broad, 
eventually terete, cortex splitting between and below subfoliar ridges; internodes 
0-5-4(-6*) mm. Leaves sessile or rarely narrowing to a pseudopetiole up to 2 mm 
long, ascending to subimbricate, sometimes tetrastichous, deciduous above base (or 
pseudopetiole) after twisting spirally; lamina 7-15 x l-l-4(-4*) mm, very narrowly 
elliptic-oblong to linear incurved to canaliculate or rarely subconduplicate, ± 
markedly cucuUate, midrib impressed beneath, margin distinct, narrowly to rather 
broader hyaline, concolorous, lucent beneath, undulate-subpapillose above, 
glaucous or not, coriaceous to somewhat succulent; apex acute, base angustate to 
parallel-sided, not or scarcely sheathing, pairs united to form narrow interfoliar 




Map 24 58. H. andinum ▲. 



See note on p. 99. 



THE GENUS HYPERICUM L. 

ridge; basal vein 1, unbranched; laminar glands rather sparse, usually visible above 
only but sometimes biseriate beneath. Inflorescence l(3)-flowered, with pseudo- 
dichotomous branches from node below and sometimes short, often congested, 
lateral branches (racemiform); pedicel absent or up to 4 mm long; upper leaves not 
transitional. Flowers 10-16 mm in diam., stellate. Sepals (5-)6-8 x 0-5-1-5 mm, 
oblanceolate-oblong to oblong or rarely linear, unequal to subequal, acute, cucul- 
late, margin ± broadly hyaline; veins 3-5, unbranched or outer branched, all not or 
slightly prominent; glands linear, punctiform in upper third. Petals yellow, 8-11 x 
2-5-4 mm, c. 1-5 x sepals, narrowly obovate-oblong; apiculus acute to acuminate; 
glands linear, distally punctiform. Stamens 25-40, longest 3-5-5 mm long, c. 0-5 x 
petals. Ovary 1-5-2-5 x 0-8-1-5 mm, ellipsoid to subglobose; styles 3(4), l-2(-3*) 
mm long, 0-5-0-9(-l-l*) x ovary, outcurved to suberect; stigmas broadly capitate. 
Capsule 4-5 x 1-5-2-5 mm, ovoid-ellipsoid to cylindric-eUipsoid, shorter than 
sepals. Seeds 0-8-1 mm long, ecarinate; testa finely scalariform. 

In damp (sometimes marshy) or sheltered areas of the open paramo; 2870-4700 m. 

Western Venezuela (Merida), Colombia (Santander to Nariiio) and northern 
Ecuador (Carchi to Cotopaxi). Map 25. 

H. lancioides differs from H. ruscoides especially in its narrower, 1 -veined, sessile 
leaves, narrower sepals and shorter styles. One collection from Colombia, Putumayo 
(Schultes & Villareal 7821), has relatively broad, petiolate leaves and longer styles 
(see characters marked*), and Cuatrecasas 17861 (Valle, Los Farallones, cerro La 
Torre) is even closer to H. ruscoides and could be of hybrid origin. 

H. lancioides is variable and comprises two subspecies, one (subsp. lancioides) 
with its most primitive form in southern Colombia and extending from there north to 
Cundinamarca and south to Cotopaxi, has a simply pseudo-dichotomous inflores- 
cence, whereas the other (subsp. congestiflorum) occurs from Cauca north-east to 
Merida and has lateral flowers crowded below the terminal flower. Intermediate 
specimens between the subspecies occur in Cauca. 



99 






59a. Hypericum lancioides Cuatrec. subsp. lancioides 

H. struthiolifolium var. genuinum Triana & Planchon in Annls Sci. nat. (Bot.) IV, 18: 
293 (1862) ['struthiolaefolium'] pro parte, quoad spec. cit. 




Map 25 59. H. lancioides a. subsp. lancioides A , h. subsp. congestiflorum A . 



100 



NORMAN K. B. ROBSON 




Fig. 14 A. H. lancioides subsp. lancioides: (a) habit; (b) leaf; (c) sepal; (d) petal; (e) stamens (partly cut 
away); (f) capsule. B. H. lancioides subsp. congestiflorum: (g) habit. C. H. selaginella: (h) habit; (i) stem 
with leaves; (j) leaf; (k) sepal; (b) petal; (m) stamens (partly cut away) and ovary; (n) capsule (a, g, h x 
1/2; b-f X 3; i-m x 4). A. B. & C. Maguire 61808; B. Cuatrecasas et al. 28120; C. Barclay & Juajibioy 

7401. 



H. strictum sensu Gleason in Bull. Torrey bot. Club 56: 102 (1929) pro parte, quoad 

spec. cit. 
H. lancioides forma pygmaea Cuatrec. in sched. 

Icon: Fig. 14A. 

Shrub 0-2-1 m tall, erect, bushy, with branches strict, pseudo-dichotomous and 
lateral. Leaves shortly petiolate or sessile, usually spreading, sometimes ± tetra- 
stichous, very narrowly elliptic or oblanceolate to linear. Inflorescence branches 
pseudo-dichotomous with flowers solitary, terminal and sometimes on well- 



i 



THE GENUS HYPERICUM L. 101 

developed branches from lower nodes. Sepals oblong to linear, with veins promin- 
ent. 

3000-4100 m. 

Southern Colombia (Cundinamarca and Meta southwards) and Ecuador (Carchi to 
Cotopaxi). 

COLOMBIA. Cauca: Cordillera Central, Paramo de las Moras, 3800-3700 m, 
12.xii.l969 (fl & fr), Cuatrecasas 27419 (BM, COL, US). Cundinamarca: Cordillera 
Oriental S. of Usme, paramo de Chisaca above Laguna Grande and L. Negra, 
9-ll.xi.l958 (fl), Barclay & Juajibioy 6191 (COL, MO, NY). Huila: Municipio de 
La Plata, Inspeccidn de Policia de Santa Leticia, Paramo de La Candelaria, 2370 m, 
15.vii.l975 (fl), Diaz, Lozano & Torres 595 (COL). Narino: Province de Pasto, 
Fuguerres, 300 m, v.1853 (fl), Triana 5464 (BM, COL, K, MEXU, NY, P, W); 
Volcan El Galeras, c. 3200-3500 m, 4.vi.l946, Schultes & Villarean960 (COL, F). 
Putumayo: Pasto to Sibundoy, Paramo de San Antonio, 2790-2880 m, 13.iii.l953 (fl 
& fr), Schultes & Cabrera 18828 (COL, GH, NY, U). Valle: Cordillera Occidental, 
Los Farrallones, cerro de La Torre, La Laguna, 3500-3550 m, l.viii.l946 (fl), 
Cuatrecasas 21870 (BM, F, P, U, US). 

ECUADOR. Carchi: Tulcan to Maldonado, c. 43 km, SW. slopes of Volcan 
Chilas, 3740 m, 6.viii.l976 (fl), 0llgaard & Balslev 8549 (AAU, BM). Cotopaxi: 
Ruminahui, 3850 m, 17.U.1947 (f[&fr), Aubertde La Rue s.n. (P). Imbabura: Lago 
San Marcos, Cayambe, 3360 m, 26.xi.1961 (fl), Cazalet & Pennington 5349 (K). 
Napos: Laguna Porjuantag, ENE. of Volcan Cayambe, 3750 m, 10. ii. 1953 (fl), 
Prescott 347 (NY). Pichincha: Paso de Guamani, 35-37 km ESE. of Quito, head- 
water of Rio Papallacta, 23.V.1947 (fl), Fosberg 27609 (NY, US); Cordillera Orien- 
tal, environs of Laguna de Chiqui, 3800 m, 4.vi.l944 (fl & fr), Acosta Solis 8176 (F). 

59b. Hypericum lancioides Cuatrec. subsp. congestiflorum (Triana & Plan- 
chon) N. Robson, comb, et stat. nov. 

H. struthiolifolium var. congestiflorum Triana & Planchon [ex Weddell, Chloris 
Andina 2: 271 (1857) 'struthiolaefolium\ nomen] mAnnls Sci. nat. (Bot.) IV, 18: 
294 (1862) pro parte, excl. syn. ; Knuth in Reprium Spec. nov. Regni veg. Beih. 43: 
484 (1927) pro parte, excl. spec. Funck & Schlim 1140. Type: Venezuela, Merida, 
Sierra Nevada de Merida, 2870 m, 1842 (fl). Linden 432 (COL?, holotype; P!, 
isotype). 

Icon: Fig. 14B. 

Shrub 01-0-5 m tall, erect and bushy or decumbent and caespitose with branches 
strict, lateral (very rarely pinnate), spreading. Leaver sessile, ascending to imbricate, 
tetrastichous, linear. Inflorescence branches elongate, sympodial (from terminal 
node) and very short, lateral, with flowers solitary or in triads in a congested 
spiciform thyrse, immediately below terminal flower or with several intervening 
sterile nodes. Sepals oblong-lanceolate, with veins not or slightly prominent. 

2870-4500 m. 

Western Venezuela (Merida), north-eastern and central Colombia (Santand6r to 
Cauca). 

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, Plan de S. 
Jose, 3625 m, 9.vi.l973 (fl), C/ee/ 10051 (U). Boyaca: Paramo de Chita, cabeceras 
del Rio Casanare, Hoya Laguna Chorros Blancos, 3340 m, 24.V.1973 (fl), Cleef 
9931A (COL, U). Caldas: Paramo del Quindio, 4100-4300 m, 15-20.viii.1922 (fl), 
Pennell & Hazen 9874 (GH, K, S). Cauca: Paramo de Purace, 3500 m, 26.v. 1944 (fl), 
Killip & Lehmann 38548 (COL). Cundinamarca: Paramo de Palacio, c. 600 m al 
NW. de Mina de Cal, 3520 m, 13.V.1972 (fl), Cleef 3733 (BM, COL, U); Lagunas de 
Buitrago, 3555 m, 27.iv.1973, Cleef 95S8A (COL). Meta: Paramo de Sumapaz, 
Cerro Nevado de Sumapaz, 3615 m, 29.1.1972, Cleef 1269 (COL, U). Risaralda: 
Municipio de Pereira, alredores de la Laguna Otun, 4000-4300 m, 3.ii.l980 (fl), 
Diaz, Valencia & Jaromillo 1754 (COL). Santander: Paramo del Almorzadero, 



102 



NORMAN K. B. ROBSON 



3500-3700 m, 20 vii.1940 (fl), Cuatrecasas & Garcia Barriga 9998 (COL, F). Tolima: 
Paramo de Herves, Eresno, 3600 m, 14.vii.l939 (fl), Hanbury -Tracy 603 (K). 

VENEZUELA. Merida: Paramo de Timotes, 3200 m, 25.1.1939 (fl), Alston 6616 
(BM, NY); Quebrada de Saisay, 3880 m, 10.iv.l930, Gehringer 57 (NY, VEN). 

In Venezuela there is a tendency in subsp. congestiflorum towards regular pyramidal 
branching, shorter glandular-punctate leaves, and smaller flowers, sometimes with 4 
styles. All these characters foreshadow H. selaginella. 



60. Hypericum selaginella N. Robson, sp. nov. 

H. lancioidei subsp. congestiflorae (Triana & Planchon) N. Robson affinis, sed 
habitu nano prostratoque, ramificatione pinnato, foliis brevioribus dense imbricatis, 
flores minoribus, stylis (3)4-5 brevioribus, differt. Type: Colombia, Boyaca, Cordil- 
lera Oriental, Sierra Nevada del Cocuy, Alto Ritacuva, Station 16, c. 4400 m, 
ll-29.iv.l959 (fl & fr), Barclay & Juajibioy lAQH (COL!, holotype; MO!, NY!, 
isotypes). 

H. selaginoides N. Robson in Cleef, Veg. Pdram. Colomb. Cord. Or.: 305 (1981), 
nomen. 

Icon: Fig. 14C. 

Shrublet up to 0-15 m tall, prostrate or ascending, forming tufts up to 150 mm in 
diam., with branches pinnate, spreading. Stems orange-brown, 6-lined and com- 
pressed when young, the subfoliar ridges broad, eventually terete, cortex splitting 
between and below subfoliar ridges; internodes c. 0-5 mm long. Leaves sessile, 
densely to loosely imbricate, tetrastichous, deciduous above base or more usually 
only with the cortex after fading; lamina 4-7 x 0-5-0-8 mm, Unear, incurved- 
canaliculate, cucullate, midrib impressed beneath, margin broadly hyaline, con- 
colorous, sublucent beneath, sublucent to dull above, sometimes subglaucous, 
coriaceous; apex acute to subacute, base parallel-sided, not sheathing, pairs united 
o-^i^ to form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense 

o+^,^ above, rather sparse or absent beneath. Inflorescence 1-flowered, with pseudo- 

c^H^^r^ dichotomous branches from node below and also short congested lateral branches 

^•^A'^ (racemiform) ; pedicel absent or up to 2 mm long; upper leaves not transitional. 

Flowers c. 8 mm in diam., stellate. Sepals 3-4 x 0-8-1-2 mm, oblong to triangular- 
lanceolate (equal), acute, cucullate, margin broadly hyaline; veins 3(5?), unbran- 
ched, slightly prominent; glands mostly linear. Petals bright yellow, tinged red 
outside, 4-5-6 x 2-2-5 mm, 1-5 x sepals, obovate-oblong; apiculus very short or 
almost absent; glands linear, distally punctiform. Stamens c. 13, longest 3-4 mm 
long, c. 0-65 x petals. Ovary 1-5-2-5 x 1-5-2 mm, subglobose; styles (3)4-5, 0-8-1 
mm long, 0-4-0-5 x ovary, outcurved; stigmas broadly capitate. Capsule c. 3 x 3 
mm, cylindric-globose to globose, shorter than sepals. Seeds 0-7-0-8 mm long, 
ecarinate; testa finely scalariform. 

In dry and stony or dampish paramo or superparamo; 3300-4340 m. 

Colombia (Boyaca, Arauca, Cundinamarca), in the Sierra Nevada del Cocuy, 
Paramo de Pisva and Macizo de Sumapaz. Map 22 (p. 94). 

COLOMBIA. Arauca: Sierra Nevada del Cocuy, Cabeceras de la Quebrada, El 
Playon, Patio Bolos, 2-5 km S. of Alto La Plaza, 4340 m, 9.iii.l973 (fl & fr), Cleef 
8944 A (COL). Boyaca: Sierra Nevada del Cocuy, Valle de los Corallitos, c. 4300 m, 
31.vii.l957 (fl), Grubb, Curry & Fernandez-Perez 178 (COL, K, US); Paramo de 
Pisva, Socha to La Punta Km 61.5, 6 km alNE.de Los Pinos, 3600 m,10.vi. 1972 (fl& 
fr), Cleef 432S (BM, COL, U). Cundinamarca: Macizo de Sumapaz, Cuchilla La 
Rabona, 3900-3950 m, 17.vii.l981 (fl), Diaz etal. 2871 (COL). 

In H. selaginella the reduction trends of H. lancioides are continued. The different 
characters between it and H. lancioides subsp. congestiflorum (dwarf prostrate habit, 
smaller and more densely imbricate leaves, smaller flowers, shorter and usually more 
numerous styles) are clear-cut except for a few collections of the latter taxon from 
Merida, which are somewhat intermediate in habit and size of parts. 



THE GENUS HYPERICUM L. 

61. Hypericum cymobrathys N. Robson, sp. nov. 

H. terrae-firmae Sprague & Riley affinis, sed habitu humiliori, foliis brevioribus 
crassioribus, interdum leviter cucullatis, supra basin deciduis, floribus minoribus in 
cymes 3-12-floris dispositis, 3-stylis, inter alia differt. Type: Colombia, Boyaca, La 
Uvita, by road from Chita, 3000-3050 m, 16. ix. 1969 (fl), Cuatrecasas & Rodriguez 
27805 (COL!, holotype; BM!, US!, isotypes). 

Shrub 0-5-1 m tall, erect, with branches ± strict, pseudo-dichotomous or occasional- 
ly lateral. Stems orange-brown to purplish-brown, 4-lined and markedly ancipitous 
when young, soon 4-angled, cortex exfoliating in irregular flakes; internodes 2-5 mm 
long. Leaves sessile, spreading from above base, tetrastichous, deciduous from 
above base without fading; lamina 9-18 x 3-8 mm, oblong to elliptic, plane, slightly 
cucullate, midrib prominent towards the base beneath, concolorous, glaucous, 
coriaceous; apex acute to obtuse, base cuneate to angustate, sheathing, saccate, 
pairs united to form narrow interfoliar ridge; basal veins 5-7, subparallel, apparently 
unbranched, tertiary reticulum not visible; laminar glands dense, prominent to ± 
impressed. Inflorescence 3-12-flowered, crowded (monochasial)?, with pseudo- 
dichotomous branches from node below; peduncle and pedicels 3-5 mm long, 
sometimes incrassate upwards; upper leaves intermediate, bracts narrowly elliptic- 
oblong, acute. Flowers c. 15 mm in diam., stellate or cyathiform. Sepals 6-8 x 
2-2-2-4 mm, narrowly lanceolate, acute; veins 5(7), unbranched, with midrib slightly 
prominent; glands linear, distally striiform. Petals bright yellow, 8-9 x 3-5 mm, c. 
1-2 X sepals, narrowly obovate; apiculus acute; glands linear, interrupted distally. 
Stamens c. 100, longest 4-5 mm long, c. 0-5 x petals. Ovary 2x1-5 mm, broadly 
ellipsoid; styles 3, 3 mm long, 1-5 x ovary, outcurving; stigmas narrow. Capsule c. 5 
X 3 mm, broadly ellipsoid, shorter than sepals. Seeds 1-2-1 -3 mm long, carinate; 
testa finely scalariform. 

Shrubby woodland on dry slopes; 2970-3500 (-3900?) m. 

Colombia (Boyaca). Apparently confined to a small area of the Cordillera Oriental 
around Guantiva and Chita. Map 11 (p. 54). 

COLOMBIA. Boyaca: Socha to Los Pinos, Km 41, 2970 m, 22.V.1973 (fl), Cleef 
9871 (BM, U); Paramo de Guantiva, 3000-3900 m, 23.vii.1952 (fl), Schneider 1225 
(S); Municipio de Tota, Cerro los Arcos, alredores de Lago de Tota, 3050 m, i.l976 
(fr), Aguirre & Rangel 391 (COL). 

Although the cymose inflorescence of H. cymobrathys is reminiscent of H. mexica- 
num, its flowers and leaves are very different. Its relationships appear to be with H. 
terrae-firmae, although the much smaller flowers, 3-12-flowered inflorescence, and 
smaller leaves make it superficially unlike it. Indeed, it appears to be an isolated 
relict species at the base of a clade that includes H. chamaemyrtus, H. gnidioides, and 
the H. gentianoides group, as well as H. denticulatum and its relatives (see part 8). 



103 




W. 




3. Acknowledgements 

I am especially grateful to Dr Bassett Maguire (NY), who has helped my studies of tropical American 
Hypericum in many ways; I have also received valuable assistance from Dr Antoine Cleef (U), Dr Jose 
Cuatrecasas (US), Dr Richard Howard (A), Dr Alicia Lourteig (P), Dr James Luteyn (NY), Dr Benjamin 
011gaard (AAU), Dr Peter Raven (MO), Dr Peter Stevens (A, GH), Dr Julian Steyermark (VEN), Dr 
Robert Wilbur (DUKE), and John Ironside Wood (British Council, Bogota). My thanks are also due to 
the directors of the following herbaria for the loan of specimens: Arnold Arboretum, Harvard University 
(A); Herbarium Jutlandicum, Aarhus (AAU); Botanical Museum, Copenhagen (C); Instituto de Ciencias 
Naturales de la Universidad Nacional, Bogota (COL); Duke University, Durham, North Carolina 
(DUKE); Botanical Museum Goteborg (GB); Gray Herbarium, Harvard University (GH); Royal Botanic 
Gardens, Kew (K); Missouri Botanical Garden, St. Louis (MO); New York Botanical Garden (NY); 
Museum National d'Histoire Naturelle, Paris (P); Naturhistoriska Riksmuseet, Stockholm (S); Institute 
for Systematic Botany, University of Utrecht (U); Instituto Botanico, Caracas (VEN); Naturhistorisches 
Museum, Vienna (W). I must also thank Miss Joan Malcolm for help with plotting distributions, and Dr 
Richard Pohl (ISC) and Dr Chris Humphries (BM) for providing photographs. 
With regard to the production of this paper, I am again deeply indebted to Mrs Margaret Tebbs for 



104 



NORMAN K. B. ROBSON 



drawing the plates and figures; I am also very grateful to Miss Maria Duda for typing most of the 
manuscript. Finally I would like to thank various colleagues for helpful discussions and suggestions, in 
particular Arthur Chater, who commented on the typescript, and Bob Press for improving my marginal 
illustrations. 



4. References 

Cleef, A. M. 1981. The vegetation of the paramos of the Colombian Cordillera Oriental. Thesis, University 

of Utrecht. 
Cuatrecasas, J. 1936. Resumen de mi actuaci6n en Colombia con motivo del II centenario del nacimiento 

de Mutis. Trab. Mus. nac. Cienc. nat., Madr. (Bot.) 33: 83-85. 
Gleason, H. A. 1929. Hypericum, subsect. Eubrathys, in northwestern South America. Bull. Torrey bot. 

Club 56: 100-107. 
Howarth, M. K. 1981. Palaeogeography of the Mesozoic. In L. R. M. Cocks (Ed.), The evolving Earth: 

197-220. London. 
Kimura, Y. 1951. Hypericaceae. In T. Nakai & M. Honda, Nova flora japonica 10. Tokyo. 
Lozano-C, G. & Schnetter, R. 1976. Estudios ecologicos en el Paramo de Cruz Verde, Colombia. II. Las 

comunidades vegetales. Caldasia 11 (54): 53-68. 
Melville, R. 1981. Vicarious plant distributions and the palaeography of the Pacific region. In G. Nelson & 

D. E. Rosen (Eds), Vicariance biogeography : a critique. New York. 
Owen, H. G. 1976. Continental displacement and expansion of the earth during the Mesozoic and 

Cenozoic. Phil. Trans. R. Soc. A, 281: 223-292. 
Raven, P. H. & Axelrod, D. J. 1974. Angiosperm biogeography and past continental movements. Ann. 

Mo. bot. Gdn 61: 529-673. 
Robson, N. K. B. 1977. Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric classification. Bull. 

Br. Mus. nat. Hist. (Bot.) 5: 291-355. 
1981 . Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the genus. Bull. Br. Mus. nat. 

Hist. (Bot.) 8: 55-226. 

1985. Studies in the genus Hypericum L. (Guttiferae). 3. Sections 1. Campylosporus to 6a. 



Umbraculoides. Bull. Br. Mus. nat. Hist. (Bot.) 12: 163-325. 
Rodriguez Jimenez, C. 1980. Hipericaceas. In R. Reitz, Flora illustrada catarinense. Itajai, Santa Catarina. 
Rosen, D. E. 1976. A vicariance model of Caribbean biogeography. Syst. Zool. 24: 431-464. 
1985. Geological hierarchies and biogeographical congruence in the Caribbean. Ann. Mo. bot. Gdn. 

72: 636-659. 
Standley, P. C. & Williams, L. 0. 1961. Flora of Guatemala, Guttiferae. Fieldiana Bot. 24 (7): 36-61. 



5. Systematic index 

Accepted names are in roman and synonyms in italic; new names are in bold, as are principal references. 
An asterisk (*) denotes a figure. 



Brathys acerosa (Kunth) Spach 

(= 20) 47 
acicularis (Kunth) Spach (= 35) 

67 
caracasana (Willd.) Spach 

(= 51a) 89 
juniperina (Kunth) Spach (-41) 

73 
juniperina L. f. (= 41) 73 
laricifolia (Juss.) Spach (= 20) 

47 
mutisiana (Kunth) Spach (= 26) 

56 
struthiolifolia (Juss.) Spach 

(= 57)96 
thuyoides (Kunth) Spach {= 13) 

37 
Hypericum sect. 29. Brathys (Mutis 

ex L.f.) Choisy 12 



acerosum Kunth (= 20) 47 
aciculare Kunth (35) 66*, 68* 
acostanum Steyerm. ex N. 

Robson (5) 27* 
andinum Gleason (58) 97, 98* 
bolivaricum N. Robson (40) 72, 

74* 
brathys Sm.{= 41)73 
brathys sensu Trev. pro parte 

(= 48b) 85 
brathys var. juniperinum (Kunth) 

Choisy (- 41) 73 
bryoides Gleason (39) 71* 
caracasanum Willd. (51) 89 
caracasanum sensu Turcz. pro 

parte (= 50) 87 
caracasanum subsp. cardonae 

(Cuatrec.) N. Robson (= 50) 

87 



caracasanum subsp. caracasanum 

(51a) 87*, 89, 91* 
caracasanum subsp. 

turumiquirense (Steyerm.) N. 

Robson (51b) 87*, 90 
caracasanum var. ocanense R. 

Keller (= 31)60 
caracasanum \dir.scherzeri{= 7) 

30 
caracasanum var. turumiquirense 

Steyerm. (- 51b) 90 
cardonae Cuatrec. (50) 87* 
carinosum R. Keller (9) 33, 37* 
cassiopiforme N. Robson (29) 

57*, 59 
chamaemyrtus sensu Gleason 

(=4)26 
costaricense N. Robson (38) 70. 

71* 



THE GENUS HYPERICUM L. 



105 



cuatrecasii Gleason (16) 39*, 41 
cymobrathys N. Robson (61) 

54*, 103 
decandrum Turcz. (30) 54*, 59, 

61* 
decorticans Planchon & Linden 

(= 8)32 
ekmanii A. H. Liogier (52) 90, 

91*, 92* 
ericifolium Steyerm. (= 4) 26 
genistoides Kunth {- 13)37 
garciae Pierce (4) 26, 27* 
gleasonii N. Robson (25) 46*, 55 
gnidioides var. polytrichoides R. 

Keller (= 30)59 
goyanesii Cuatrec. (14) 38, 39* 
graciliforme N. Robson (= 48b) 

85 
gyropodioides sensu Jackson & 

Hooker (= 12)36 
hartwegii Bentham (22) 46*, 51 
hartwegii sensu Triana & 

Planchon pro parte (= 19)45 
hartwegii var. patens R. Keller 

(= 13)37 
hilaireanum ( = 6) 28 
holtonii Gleason (= 41) 73 
horizontale N. Robson (46) 81, 

82* 
incurvum Urban (= 2a) 21 
irazuense Kuntze ex N. Robson 

(7) 30, 31*, 32* 
jahnii R. Keller {= 41)73 
jaramilloi N. Robson (49) 86, 87* 
juniperinum Kunth (41) 72, 74* 
lancifolium Gleason (45) 77*, 80 
lancioides Cuatrec. (59) 98 
/andojd^5 sensu Steyerm. (= 45) 

80 
lancioides subsp. congestiflorum 

(Triana & Planchon) N. 

Robson (59b) 99*, 100*, 101 
lancioides subsp. lancioides (59a) 

99*, 100* 
lancioides forma pygmaea 

Cuatrec. (59a) 100 
laricifolium Juss. (20) 47, 48*, 

49* 
laricifolium sensu Bentham 

(=35)67 
laricifolium var. acerosum 

(Kunth) Weddell (= 20) 47 
laricifolium var. glaucum R. 

Keller (= 48b) 85 
laricoides Gleason (= 20) 47 
lindeniiR. Keller {= 19)45 
lindenii sensu Cuatrec. (= 15) 40 
llanganaticum N. Robson (56) 

95, 96* 
loxense Bentham (18) 43 
loxense subsp. aequatoriaie (R. 

Keller) N. Robson (18a) 44* 
loxense var. aequatoriaie R. 

Keller (= 18a) 44 
loxense subsp. loxense (18b) 44*, 

45 



lycopodioides Triana & Plauchon 

(12)36 
magdalenicum N. Robson (32) 

62*, 63 
magniflorum Cuatrec. (24) 53*, 

54* 
maguirei N. Robson (23) 46*, 52 
marahuacanum N. Robson (44) 

77 
marahuacanum subsp. 

chimantaicum N. Robson (44c) 

71*, 78*, 80 
marahuacanum subsp. 

marahuacanum (44a) 71*, 78*, 

79 
marahuacanum subsp. 

strictissimum N. Robson (44b) 

71*, 78*, 79 
martense N. Robson (21) 48*, 

49*, 51 
meridense Steyerm. (= 8) 32 
mexicanum L. (26) 56, 57* 
? mexicanum sensu Choisy 

(= 26) 56 
mexicanum var. intermedium 

Kuntze (=26) 56 
millefolium Urban & Ekman (54) 

91*, 92*, 93 
mutisianum Kunth (=26) 56 
myricariifolium Hieron. (15) 40, 

44* 
myricariifolium Hieron. ex R. 

Keller (= 15)40 
papillosum N. Robson (11) 24*, 

34, 35* 
parallelum N. Robson (43) 76, 

77* 
phellos Gleason (3) 23, 24* 
phellos subsp. oroqueanum N. 

Robson (3b) 25 
phellos subsp. phellos (3a) 23 
phellos subsp. platyphyllum 

(Gleason) N. Robson (3c) 25 
pimeleoides Planchon & Linden 

ex Triana & Planchon (31) 60, 

62* 
piriai (6) 28, 29* 
platypetalumTurcz. (= 20)47 
platyphyllum Gleason (= 3c) 25 
platyphyllum Gleason pro parte 

(= 25)55 
prietoi N. Robson (28) 57*, 58 
prostratum Cuatrec. (42) 75, 77* 
pseudobrathys Turcs. (— 41)73 
pseudobrathys Turcs. pro parte 

(= 48b) 85 
pycnophyllum Urban (53) 91*, 

92* 
quitense R. Keller (17) 39*, 41, 

42* 
racemulosum Turcz. (= 20) 47 
recurvum N. Robson (36) 68* 
resinosum Bentham (= 26) 56 
rimbachianum Dieh {= 17)41 
ruscoides Cuatrec. (55) 93, 

94* 



sabiniforme Trev. (19) 45, 46* 
selaginella N. Robson (60) 94*, 

100*, 102 
selaginoides N. Robson (= 60) 

102 
silenoides sensu R. Keller (=7) 

30 
simonsii N. Robson (10) 24*, 34, 

35* 
sprucei N. Robson (34) 62*, 65, 

66* 
stenoclados Cuatrec. (= 9) 33 
stenopetalum Turcz. (8) 31*, 32* 
stenopetalum var. majus Triana 

& Planchon (= 10) 34 
strictum Kunth (48) 83 
strictum sensu Gleason pro parte 

(= 59a) 100 
strictum subsp. compactum 

(Triana & Planchon) N. 

Robson (59b) 85* 
strictum subsp. strictum (48a) 84, 

85* 
strictum var. compactum (Triana 

& Planchon) Gleason (= 48b) 

85 
strictum var. reductum Gleason 

(= 41)73 
struthiolifolium Juss. (57) 96* 
struthiolifolium Juss. pro parte 

(= 35)67 
struthiolifolium sensu Knuth 

(= 45)80 
struthiolifolium sensu R. Keller 

pro parte (= 34) 65 
struthiolifolium sensu Trev. pro 

parte (= 9)33 
struthiolifolium var. compactum 

Triana & Planchon (= 48b) 85 
struthliolifolium var. 

congestiflorum Triana & 

Planchon (= 59b) 101 
struthiolifolium var. 

congestiflorum sensu Kunth 

pro parte (= 41) 73 
struthiolifolium var. genuinum 

Triana & Planchon (= 57) 96 
struthiolifolium var. genuinum 

Triana & Planchon pro parte 

(= 59a) 99 
struthiolifolium var. gracile 

Triana & Planchon (= 48b) 85 
struthiolifolium var. minutum 

Choisy (= 35)67 
struthiolifolium var. parviflorum 

R. Keller (= 58)97 
struthiolifolium var. P sensu 

Choisy pro parte (= 35) 67 
struthiolifolum var. strictum 

(Kunth) Choisy (= 48a) 84 
stuebelii Hieron. (27) 54*, 58 
stuebelii Hieron. ex R. Keller 

(= 27) 58 
styphelioides A. Richard (2) 20 
styphelioides sensu Standley & 

Williams (= 1) 18 



106 



NORMAN K. B. ROBSON 



styphelioides subsp. clarense 

Lippold (2a) 21* 
styphelioides subsp. moaense 

Lippold (2c) 21*, 22 
styphelioides subsp. 

styphelioides (2b) 21* 
tachirense Steyerm. (= 3a) 23 
tamanum Cuatrec. (= 3a) 23 
tenuifolium St.-Hil. (= 6) 28 



terrae-firmae Sprague & Riley 

(1)18, 19*, 21* 
tetrastichum Cuatrec. (47) 82* 
thuyoides Kunth (13) 37* 
thymifolium Cuatrec. (= 3a) 23 
thymifolium sensu Planchon & 

Triana pro parte (= 9) 33 
r/jyAnZ/b/ZuAM sensu Turcz. (= 19) 

45 



trianae N. Robson (= 19) 45 
valleanum N. Robson (33) 62*, 

64,66* 
weberbaueri R. Keller (= 57) 96 
weberbaueri sensu Gleason 

(= 45)80 
wurdackii N. Robson (37) 62*, 

69 



Studies in the genus Hypericum L. (Guttiferae) 

N. K. B. Robson 

When completed in nine parts, this series of papers will constitute a monograph of 
Hypericum, St John's wort. This ancient genus, with an almost world-wide distribu- 
tion, is of great scientific interest, as well as of considerable importance in horticul- 
ture and, to a lesser extent, pharmacy. Parts 1 (1977) and 2 (1981) are introductory, 
and the detailed taxonomy begins with part 3 (1985), in which the first seven sections 
(sects l-6a) are treated. The present paper (part 7) contains the majority of sect. 29, 
the remainder of which will be included in part 8, along with accounts of sects 30 and 
31. Subsequent papers (parts 4-6) will contain treatments of sects 7-28, and the 
concluding paper (part 9) will contain addenda, corrigenda, a revised sectional key 
and enumeration, and a continuous numerical series of species. 

Bulletin British Museum (Natural History): Botany Series 

Vol. 5 No. 6 Studies in the genus Hypericum L. (Guttiferae). 1. Infrageneric 
classification. N. K. B. Robson. 1977. Pp. 291-355, 9 figs. £7.25 

Vol. 8 No. 2 Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the 
genus. N. K. B. Robson. 1981. Pp. 52-226, 73 figs, 8 tables. £23.50 

Vol. 12 No. 4 Studies in the genus Hypericum L. (Guttiferae). 3. Sections 1. 
Campylosporus to 6a. Umbraculoides . N. K. B. Robson. 1985. Pp. 163-325, 24 
figs, 34 maps. 565 08003 2. £28.80 



Titles to be published in Volume 16 



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

ByN. K. B.Robson 

The lichen genus Ramalina in Australia 

By G. N. Stevens 

An annotated list of vascular plants collected in the valleys south of 
Mt Everest 

By G. Miehe 

An illustrated catalogue of type specimens in the Greville diatom herbarium 

By D.M.Williams 



Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk 
Printed in Great Britain by Henry Ling Ltd, Dorchester 



\^ - ~ - J ■V—' I 



Bulletin of the i 

British Museum (Natural History) 



The lichen genus Ramalina in AustraUa 
G. Nell Stevens 



Botany series Vol 16 No 2 25 June 1987 



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© British Museum (Natural History), 1987 



The Botany series is edited in the Museum's Department of Botany 
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Botany series 

Vol 16 No 2 pp 107-223 



Issued 25 June 1987 



The lichen genus Ramalina in Australia 

i! PR. 

G. Nell Stevens 

Department of Botany, University of Queensland, St Lucia, 4067, Australia 

Contents 

Synopsis 108 

Introduction 108 

The Australian environment 110 

Historical understanding of Ramalina and Ramalinaceae 1 15 

Collectors of Australian Ramalinae 117 

Materials and methods 120 

Results 121 

Morphology 121 

Anatomy 126 

Chemistry 132 

Correlation between secondary-product chemistry and ecogeography 

in the Ramalina subfraxinea complex 134 

Distribution patterns in relation to climatic indices 135 

Biogeography 140 

Palaeobiogeography 142 

Taxonomy 147 

Taxonomic ranks 147 

Hybridization 148 

Description of the genus Ramalina 149 

Enumeration of the Australian taxa 150 

Key to Ramalina in Australia 150 

1. R. aUstr aliens is 152 

2. R. caespitella 155 

3. R. canariensis 156 

4. R. exiguella 158 

5 . R. filicaulis 160 

6. R. fimbriata 161 

7. R. glaucescens 163 

8. R.litorea 169 

9. R. peruviana 170 

10. R.reducta 175 

11. R.tenella 176 

12. R. unilateralis 177 

13. R. whinrayi 179 

14. R.celastri 180 

15. R.inflata 185 

16. R.fissa 193 

17. R. nervulosa 195 

18. R.pacifica 201 

19. R. subfraxinea 203 

20. R. tropica 210 

Appendix. Disposition of exluded Ramalina taxa 217 

Acknowledgements 217 

References 218 

Index 222 



Bull. Br. Mus. nat. Hist. (Bot.) 16 (2): 107-223 Issued 25 June 1987 



108 G. NELL STEVENS 

Synopsis 

This paper presents a revision of the lichen genus Ramalina in AustraUa. Of the 60 Ramalina taxa 
previously listed for Australia in catalogues only nine of those names are retained, six with unchanged 
status. Twenty-eight taxa belonging to 20 species are recognized for Australia. Two are new species: R. 
filicaulis and R. tropica. A new variety R. subfraxinea var. norstictica and a new subspecies R. inflata subsp. 
australis are described, and four names, R. celastri subsp. ovalis, R. inflata subsp. perpusilla, R. subfraxinea 
var. confirmata, and R. subfraxinea var. leiodea are new combinations. 

The taxonomically important features of this genus (viz. morphology, anatomy, and chemistry) are 
discussed. In delimiting taxa, the morphological plasticity shown by this genus has been taken into 
consideration so that a wide range in infraspecific variation has been accepted for some of the taxa. The 
distribution patterns of the Australian Ramalinae are shown to closely follow the distribution of Australian 
phanerogam groups, which are controlled by the environmental parameters of rainfall and temperature; 
this enables the Ramalinae to be conveniently divided into three response groups (viz. megatherm, 
mesotherm, and micro therm) as used to divide the AustraUan phanerogams. On the basis of species 
diversity it is hypothesised that the genus had its origin in west Gondwanaland (South America and Africa) 
and then spread to the east. It is suggested that subsequent speciation occurred within Australia. 

Introduction 

The genus Ramalina has been studied in the northern hemisphere for over two hundred years 
and several hundred taxa have been named. In the southern hemisphere very little research has 
been undertaken and no revisional work had been carried out on the Australian Ramalinae 
before the present study. During the course of previous research into the taxonomy and ecology 
of twenty-four lichen genera found on eastern Australian mangroves (Stevens, 1978) it became 
apparent how necessary it was for a revision of the Australian representatives of the genus to be 
undertaken. 

This present work provides a taxonomic revision of the Australian Ramalinae as well as an 
ecological survey and a biogeographic assessment of the genus. Research into the Australian 
taxa in this genus also contributes to the overall understanding of the biogeography of allied 
Ramalina species which occur in the other Gondwanaland regions of Africa, India, and South 
America. 

The genus is regarded as a difficult one and its morphological plasticity is renowned. Howe 
(1913-14) aptly summed up the dilemma when he stated 'there is little difficulty in recognizing 
the characteristics of the genus Ramalina. The species on the other hand, have presented a most 
difficult problem, caused largely no doubt by the innumerable intergrades that present them- 
selves, and by the unwarranted description of new species'. 

This problem was encountered by the author when confronted by the record of 60 different 
Ramalina taxa reported for Australia (Wetmore, 1963; Weber & Wetmore, 1972) in catalogues 
which listed all taxa recorded for Australia in papers published from 1804 to 1972. It was 
apparent quite early in the study that many of these names were incorrect; some were synonyms, 
others misapplication of the names of northern hemisphere species which they superficially 
resembled. 

Other recent mention of Australian Ramalinae occurs in an unpublished key to the genus 
Ramalina by Dahl (1971) and in Filson & Rogers (1979), but in both of these accounts little 
preliminary research had been carried out before naming the taxa. Most of the existing literature 
summarizing Australian material was therefore of little help in the present study. 

There are few modern publications recording research which include the southern hemi- 
sphere Ramalinae. Krog & Swinscow (1974, 1975, 1976) have published detailed work on the 
East African Ramalinae. Rundel (1978fl) has dealt with the Ramalina usnea complex in North 
and South America and in a paper on the lichens of Tristan da Cunha, J0rgensen (1977) 
discussed six species oi Ramalina. No other publications discuss this genus in detail; however, a 
series of papers by Osorio published from 1967 to 1983 list the lichens found in Uruguay, 
Paraguay, Brazil, and Argentina and these include some species oi Ramalina. An account of the 
lichens of New Zealand (Galloway, 1985) which includes the genus Ramalina lists only nine 
species. 



RAMALINA IN AUSTRALIA 



109 



During the present research four months were spent in Europe and U.S.A. visiting herbaria 
(BM, FH, H, HAW, O, S, UPS and US) and becoming famiUar with northern hemisphere 
Ramalina species. This enabled an insight to be gained into the reason why AustraUan species 
were often given the names of the northern hemisphere species. The type material of species 
which occur in other southern continents was also examined where possible. 

In this study the approach has been to reinforce the taxonomic revision of the genus by 
incorporating ecological data in order to explain phenotypic variation, and to enable the 
delimitation of the taxa to be made more confidently. The extreme morphological variation 
within the species was a major problem. The extent of such variation in Australian material had 
not previously been gauged, nor was there any conception of the evolutionary relationships that 
might exist between the different taxa. 

Many thousands of specimens therefore needed to be examined before any decisions could be 
made on the delimitation of species. Examination of too few specimens makes it impossible to 
understand the breadth of morphological plasticity exhibited by a single species, and the early 
taxonomy of Australian Ramalinae was based on single or at most several thalli which were 
usually collected by someone other than the taxonomist. 

Collection of Ramalina material by the author began in 1975, when the lichens on mangroves 
were studied (Stevens, 1978). Since then collecting trips have covered parts of New South 
Wales, Victoria, Tasmania, Western AustraUa, and Northe i Territory and innumerable 
collecting forays were made in many areas of Queensland. After several expeditions inland it 
was found that the genus Ramalina was restricted to a zone varying within 100-200 km of the 
coastline, thus eliminating the need to collect in the arid interior (Fig. 1). 

Observation of the different ecological habitats occupied by any one species was made to 
enable an appreciation of the variation that occurs in the morphology of a species under different 
environmental pressures. As well as sampUng different habitats in the one region, thalU were 




Fig. 1 Ramalina distribution in Australia. 



110 G. NELL STEVENS 

collected (wherever possible) from latitude to latitude in order to assess if a continuum existed 
between the different morphologies found over a wide geographical range. 

To a large extent this research has been carried out on personal collections of fresh material, 
supplemented by collections made by present day Australian lichenologists, material available 
from herbaria in each of the Australian states, and to a far lesser extent on old collections held in 
overseas herbaria. The latter proved to be of httle use as only vague information concerning the 
location of the collection was ever recorded, e.g. New Holland. In contrast the Australian 
material was well documented on the whole but the specimens remained unnamed or bore 
doubtful identifications. 

All of the AustraUan types were examined and, where possible, compared with fresh material 
from the type location. In this way the amount of variation that occurred within the particular 
species could be assessed. It was found in some instances that the type represented one extreme 
in a large population of morphotypes. 

An attempt has been made to show possible evolutionary relationships between some of the 
taxa, based on morphological and/or chemical evidence associated with geographical patterns of 
distribution, as suggested by Imshaug & Brodo (1966). 

This revision has treated all Australian material as belonging to the genus Ramalina Ach. , as 
observations during the present study did not substantiate separation into the two genera 
Ramalina Ach. and Fistulariella Bowler & Rundel. No species belonging to any other divisions 
within the Ramalinaceae (Bowler, 1981) occur in Australia. A representative collection has 
been deposited in BM. 

The Australian environment 

Although it is beyond the scope of this study to provide a detailed discussion of the geography, 
cUmate, botany, and geology of Australia, a general outline is presented to enable a better 
appreciation of the factors which are influencing Ramalina distribution (Fig. 1). 

(1) Geography 

The area of Australia is approximately 8,700,000 km^, larger than all of western Europe, and its 
immense coastline measures some 36,700 km. In total it comprises the AustraUan mainland, the 
large island of Tasmania, and numerous small islands around its coastline. It is washed by three 
oceans, the Pacific, Southern, and Indian, and four seas, the Tasman, Timor, Coral, and 
Arafura. It extends from approximately 9°30'S (Dauan Island in Torres Strait) to 43°S (southern 
Tasmania) and includes Macquarie Island at 55°S; its climate varies from equatorial through 
subtropical in the north to warm temperate, cool temperate to sub-antarctic in the south. 

Overall the landmass is low: exceeding 1000 m in only very limited areas, most of these being 
close to the eastern coastline, predominantly in a north-south alignment through Queensland, 
New South Wales, and Victoria. This continuation of the Eastern Highlands and Great Dividing 
Range from north to south creates a fairly uniform topography which is conducive to species 
dispersal over a wide geographical area without creating any substantial barrier to the distri- 
bution, which could lead to endemism. 

In the south this mountain range turns west through the Victorian Alps to the Grampians of 
western Victoria. Farther south the highland chain is apparent as islands in Bass Strait and in 
Tasmania. During times of low sea-level in the Pleistocene glacial period, these islands and 
Tasmania were joined to the mainland. Two endemic species have evolved, probably as a result 
of isolation of these islands from the continent. Several well separated parts of the mountain 
chain have individual mountains which exceed 1500 m elevation. These are found in Tasmania, 
the Victorian Alps, the Snowy Mountains, the New England Tableland in New South Wales, 
and Bellenden Ker Range in Queensland. 

The majority of the western and southern AustraUan land mass is a vast flat peneplain with an 
elevation between 200 m and 500 m. In the north-west area the highest peak in the Hamersley 
Ranges exceeds 1200 m in height, and in central Australia the Macdonnell and Musgrave Ranges 
reach a similar height. Much of the interior of Australia is arid sand-ridge desert; the arid zone 



RAMALINA IN AUSTRALIA 111 

extends to the western and southern coasts between the moist south-west corner of Western 
AustraUa and the more fertile south-east of Australia. Extensive salt lakes occur in the south and 
south-western regions of the arid interior. 

(2) Climate 

The general cHmatic characteristics of Australia are dictated by its latitudinal position lying 
astride the mid-latitude high pressure belt. The prevaiUng wind patterns and general aridity over 
continental Australia reflect the dominating control of eastward-moving anticyclonic cells which 
track between 37°-38°S in summer and between 29°-32°S in winter. Seasonal change is linked 
with these sun-controlled shifts in the paths of the anticyclonic cells and associated movements 
of the south-easterlies and the inter-tropical convergence zone which influence the climate of 
eastern and northern Australia, and the westerlies which influence the climate of southern 
AustraUa and Tasmania (Nix, 1981). 

Summer rainfall predominates in the north, winter rainfall in the south, and along the eastern 
coast a more uniform distribution of rainfall is common (Figs 2A & B). The eastern coastline of 
Australia receives a substantial rainfall, which ranges from over 2000 mm in north Queensland 
to 741 mm in Westernport Bay, Victoria. The western coastline receives far less precipitation 
per annum, partly due to the lack of any mountain ranges close to the coast. The annual 
precipitation ranges between 230 mm and 840 mm per annum with the highest recordings in the 
southwest corner where winter rains occur. Winter rainfall is predominant as far north as 24°S, 
then summer rainfall becomes dominant in the monsoonal area. 

More than 80% of the continent has at least three months each year which are without 
effective precipitation (Nix, 1981). Two-thirds of Australia has less than 500 mm rainfall per 
annum and one-third less than 250 mm. Most of the coastline represents a moist rim around an 
otherwise dry continent; however parts of this coastline in South and Western Australia are also 
extremely arid, and no Ramalinae occur there. The absence of Ramalinae in the arid inland is 
possibly due to a combination of inhibiting factors such as high summer temperatures, low 
rainfall, and lack of a suitable substrate. 

In tropical Australia the highest relative humidity occurs during the rainy summer season 
while in the temperate region relative humidity is generally highest in winter and lowest during 
the summer. The relative humidity variation during the day closely follows the diurnal variation 
of temperature, being highest with low temperatures and lowest with high temperatures. 

Mean minimum temperatures occur in July throughout AustraHa. Freezing temperatures and 
frosts occur regularly below the 5°C isotherm and the frost-free limit coincides approximately 
with the 10°C isotherm. The ameliorating influence of the surrounding oceans is evident in the 
coastal areas of southern Australia and Tasmania. Rainfall is the dominant mode of precipi- 
tation, with sleet and snow confined to the higher mountains of southeastern Australia on the 
mainland and in Tasmania. 

The greater part of the AustraUan land mass receives over 3000 hours of sunshine per year 
while the east coast is reported to receive between 2500 and 3000 hours, indicating that 
cloudiness is higher near the coast. 

The Queensland coast as far south as Mackay and the Western Australian coast as far south as 
Carnarvon are subject to tropical cyclones, normally between December and March, which 
cause very heavy rainfall and accompanying strong winds. Sea surface temperatures around 
Australia range from 15°-30°C in summer to 12°-26°C in winter. 

(3) Botany 

Specht (1970) arbitrarily divided the Australian vegetation into structural formations based on 
foliage cover and life form of the upper stratum (Table 1). The structure of the community and 
the species therein reflect the amount of moisture available to the plants e.g. tall dense plant 
communities are found in high rainfall areas whereas more stunted open communities occur in 
drier regions. Edaphic factors also influence the eventual structural formation. 

The majority of Ramalina collection locations lie in those plant communities hsted on the top 
left hand side of Table 1 , representing those which occur in relatively high rainfall areas and have 



112 



G. NELL STEVENS 




Fig. 2 A. Summer seasonal rainfall (November- April) equalled or exceeded 8 years in 10; B. Winter 
seasonal rainfall (May-October) equalled or exceeded 8 years in 10 (adapted from Nix, 1981). 



RAMALINA IN AUSTRALIA 
Table 1 Structural formations in Australian plant communities (modified from Specht, 1970). 



113 



* 




Foliage projective cover of tallest stratum 




Life form of 












tallest stratum 


100-70% 


70-50% 


50-30% 


30-10% 


<10% 


Trees >30 m 


Tall cbsed- 
forest 


Tall 
open-forest 


— 


— 


— 


Trees 10-30 m 


Closed- 
forest 


Open-forest 


Open-forest 


Woodland 


Open- 
woodland 


Trees < 10 m 


Low closed- 


Low 


Low 


Low 


Low open- 




forest 


open-forest 


open-forest 


woodland 


woodland 


Shrubs >2 m 


Closed 


Open-scrub 


Open-scrub 


Tall 


Tall open- 




scrub 






shrubland 


shrubland 


Shrubs 0-25-2 m 












Heathy 


Closed- 


Open- 


Open- 


Low 


Low open- - 




heathland 


heathland 


heathland 


shrubland 


shrubland 


Chenopodiaceous 


— 


— 


Low 


Low 


Low open- 








shrubland 


shrubland 


shrubland 


Shrubs <0-25 m 


— 


— 


— 


Dwarf open- 
heathland 


Dwarf open- 
heathland 


Hummock grasses 


— 


— 


— 


Hummock 
grassland 


Open hummock 
grassland 



a relatively high Foliage Projective Cover (FPC) e.g. (1) Closed-forest, (2) Tall open-forest, (3) 
Open-forest, and (4) Open-scrub. These structural formations are made up of overstorey and 
understorey vegetation, and depending on the amount of canopy cover, vary in FPC from 100% 
to 30%. They can be briefly described as follows: 

(a) Closed-forest (also referred to as rain-forest, it can also describe some mangrove forests). 
This community comprises many different tree genera and species which alter in composition 
with change in latitude. It is found in disjunct pockets from Cape York to Tasmania and grades 
from tropical mesophyll vine forests (with leaf size 45-180 cm^) to subtropical mesophyll and 
notophyll vine forest (with leaf size 20-25-45 cm^) to warm temperate notophyll vine forest to 
cool temperate microphyll moss forest (with leaf size 2-25-20-25 cm'^). 

Closed-forest communities occur in humid areas where rainfall exceeds 1500 mm per annum; 
overstorey FPC in such a formation approaches 100%. These formations reach their best 
development on the basaltic soils of the Eastern Highlands. Several species of Ramalina grow in 
this habitat, two species are specific to the rain-forest canopy, and another three species are 
found in both closed-forest and open-forest communities. 

These closed-forests can contain trees which exceed 30 m in height, but in monsoonal 
northern Australia and coastal Queensland, low closed-forests occur which are composed of 
trees reaching less than 10 m in height. During the wet season these forests have a FPC of 100% , 
but during the dry winters the FPC may be as low as 70% as some of the trees are semi- 
deciduous. In inland Queensland the low closed-forest is the only formation which supports any 
Ramalina species. 

(b) Tall open-forest (with sclerophyllous understorey with or without tree ferns) occurs in 
sub-humid sites of eastern, south-eastern and south-western Australia. Overstorey FPC in such 
a community varies from 70% to 50%. The dominant tree species in the overstorey is 
Eucalyptus, whereas the low tree/tall shrub stratum (less than 10 m high) contains 18 genera of 
angiosperms (Specht, 1981a). Several species of Ramalina have been found growing on twigs in 
the understorey, but rarely do species of Ramalina grow on eucalypts. 

(c) Open-forest (with either heathy sclerophyll or grassy understorey). This formation is the 
most widespread in Australia, occurring in the north, east, and southern parts of the continent; 
each area experiences quite different rainfall patterns ranging from summer rains to summer- 
winter uniform rainfall, to winter rains respectively. The genus Eucalyptus dominates these 



114 G. NELL STEVENS 

communities, although species of Casuarina and Acacia are also plentiful. The overstorey FPC 
for open-forest communities varies between 70-30% . No Ramalina species grow in the tropical 
open-forests, but occur on Casuarina and Acacia in the temperate communities, 
(d) Mallee open-scrub (with savannah-chenopod understorey) . This structural formation is 
common in the drier Mediterranean climate of southern Australia where species of Eucalypt 
dominate, but other genera, including Acacia are scattered between the mallee eucalypts. 
Overstorey FPC in these areas can be as low as 30%. The relatively low rainfall (250-800 mm 
p. a.) occurs in winter, followed by hot dry summers. Four species of Ramalina have been 
collected in this formation. 

Serai plant communities exist along the coastal fringe of Australia; these are coastal dune 
communities and saline coastal communities, the latter made up of salt-marsh, mangrove 
vegetation, and marine meadow (Specht, 1981a); only the coastal dune community and the 
mangroves act as phorophytes for Ramalina species. Such communities usually produce a series 
of structural formations, influenced by micro-habitat conditions; the overstorey FPC therefore 
differs in different parts of the community. 

(A) Coastal dune community: In the tropics, lowland rain-forest abuts the open coastline in 
areas where mangroves are not dominant; in the subtropics the sand dune vegetation comprises 
various shrub species of Banksia integrifolia, Callitris columellaris , and Casuarina equisitifolia as 
the dominant tree species. Along the southern coast-line, the foredunes are vegetated by heathy 
shrubs; Pimelea serpyllifolia, Leptospermum laevigatum, Leucopogon parviflora, Monotoca 
elliptica, Bursaria spinosa, and Acacia sophorae. Several Ramalina species occur in this 
community, the species varying with change in latitude and longitude. 

(B) The mangroves (saline coastal community): This vegetation occupies the muddy areas at 
the mouths of rivers and bays along the eastern, north-western, and northern coastlines of 
Australia, with pockets in South AustraUa. The structure of such communities varies from 
closed-forest to woodland to tall shrubland. The major concentration of mangrove vegetation 
(approx. 28 species) is confined to the tropics gradually dwindUng in species numbers in the 
subtropics and eventually being reduced to one species, Avicennia marina, towards the 
temperate zone. Both FPC and the amount of rainfall vary considerably depending on the 
latitude of the mangrove community. 

Those mangrove species which act as phorophytes to the RamaUnae are Rhizophora stylosa, 
Rhizophora apiculata, Ceriops tagal, Bruguiera gymnorhiza, Avicennia marina, Aegiceras 
corniculatum, Lumnitzera racemosa, and Excoecaria agallocha. 

(4) Geology 

The saxicolous species of Ramalina are totally dependent on the presence of rock outcrops 

within their distribution range. Corticolous species, however, are only indirectly influenced 

through the presence or absence of particular plant communities which grow on the different soil 

types. 

The four main rock types involved in the areas where Ramalina species occur are (1) basaltic 
rocks (2) metamorphics and fine grained sedimentary rocks (3) sandstones and (4) granites. 

The major areas of basalt are Tertiary in age, and are confined to the Eastern Highlands. Most 
of these elevated areas are forested with either rain-forest or tall open-forest. Additional areas 
of basalt of Quaternary age are found in north-east and central-east Queensland, supporting low 
closed-forests, and western Victoria at lower altitudes on the coastal plain which supports 
extensive areas of open-woodland and grassland. 

In eastern Australia the metamorphic rocks and many fine-grained, compacted sedimentary 
rocks which are associated with them are of Palaeozoic age, and have been folded and elevated, 
forming mountain ranges and hilly country in the Eastern Highlands. They have been intruded 
by granite batholiths, which also make up large areas, having been revealed by prolonged 
erosion e.g. New England, Murrumbidgee, Kosciusko, and many other masses in the eastern 
states. These are areas of high elevation subject to cold winters. 

The sandstones which are most important are those sub-horizontal deposits of Mesozoic age 



RAMALINA IN AUSTRALIA 115 

forming for example, much of the surface of the Sydney Basin, Moreton Basin, and the 
Carnarvon Range. The Grampians are composed of dipping sandstones of Palaeozoic age. 

In South Australia and Western Australia only a few species of Ramalina occur on trees 
growing on soils derived from old granites and metamorphic rocks (schists and quartzites). 
There are very few occurrences of Ramalinae in areas of calcareous rocks of the Tertiary basins 
and Pleistocene dune deposits. 

Historical understanding of Ramalina and Ramalinaceae 

(1) THE GENUS RAMALINA 

The genus Ramalina was first described by Acharius (1810) when he distinguished it from 

Parmelia by its cartilaginous thallus and apothecial characters. 

Acharius (1810) listed ten species (together with their forms) as belonging to the genus, 
namely Ramalina homalea, R. linearis, R. complanata, R. peruviana, R. polymorpha, R. 
fraxinea, R. fastigiata, R. scopulorum, R. farinacea, and R. pollinaria. Except for the first, all of 
these species have been retained in the genus to the present day. 

Many more taxa have been added to this original number in the last 175 years, some being 
removed later into separate genera because of differences found in the anatomy, the spore shape 
and size, or the colour of pycnidia. Montagne (1852) examined the cortex oiR. homalea Ach. 
and found that the hyphae were arranged perpendicular to the surface without any mechanical 
support tissue in the cortex; he therefore erected a new genus Desmazieria Mont, to incorporate 
all Ramalina species possessing this anatomical feature, using R. homalea as the type. 

Massalongo (1854) erected the genus Cenozosia Massal. based on R. inanis Mont, as the type 
and characterized by its fistulose spongy thallus with a single layer of branching cells in the 
cortex. This monotypic genus has been retained as a distinct taxon up to the present. 

By using the colour of spermatia, as well as anatomical and thallus differences, Stizenberger 
(1862) divided the genus Ramalina into three parts (i) Cenozosia (Massal.) (type R. inanis Mont.) 
as a species with thallus hollow, spermatia black (ii) Desmazieria (Mont.) (type R. ceruchis Ach.) 
for those species with simple cortex, thallus interior cottony and spermatia black and (iii) 
Euramalina Stizenb. (type R. scopulorum Retz.) for species having a double cortex, thallus 
interior cottony, and spermatia pale or colourless. 

Nylander (1870) also used colour of spermogonia conceptacles for initial division within the 
genus, separating species with (1) totally black spermatia (2) partially black or (3) pale or 
colourless spermatia. Within group (1) he then used differences in anatomical structure of the 
cortex to separate R. inanis Mont. (= Cenozosia Massal.) from the rest. He placed five other 
species in another group which corresponds to Desmazieria Mont. , namely R. ceruchis Ach. , R. 
combeiodes Nyl. , R. homalea Ach, , R. flaccescens Nyl. , and R. testudinaria Nyl. 

Ramalina melanothrix Laurer was separated from the rest because of its two-layered cortex 
(= Euramalina Stizenb.); its present position is in the genus Trichoramalina Rundel & Bowler. 
Group (2) contained only R. carpathica Korber, a species which Bowler & Rundel (1977) placed 
in the genus Fistulariella Bowler & Rundel. Group (3) contained 56 species which today belong 
in three genera, Ramalina Ach. , Niebla Rundel & Bowler, and Fistulariella Bowler & Rundel. 

Vainio (1890) also laid stress on anatomical structure of the cortex, accepting section 
Euramalina Stizenb. as having a chondroid cortex made up of longitudinal hyphae, and sections 
Desmazieria (Mont.) and Cenozosia (Massal.) as having a cortex made up of transverse hyphae. 
Vainio divided Euramalina Stizenb. into two stirps based on thallus structure (1) Fistularia 
Vainio species with inflated hollow branches and (2) Myelopoea Vainio species with a con- 
tinuous arachnoid or cottony medulla. He used branch shape to subdivide stirps Myelopoea into 
series Teretiusculae Vainio (containing species with terete or angular branches) and series 
Compressiusculae Vainio (containing species with flattened branches rarely two-edged). 

Hue (1901) followed Vainio in this division of Ramalina when he divided the genus into three 
groups on morphology: (A) Teretiusculae Vainio (B) Compressiusculae Vainio and (C) Fistulo- 
saeHxiQ. 

Steiner (1904) based his divisions oi Ramalina on the arrangement of cortical tissue external to 



116 G. NELL STEVENS 

the algal layer. His section Corticatae (outer layer consisting of more or less transversely 
arranged conglutinated hyphae) was equivalent to Desmazieria Mont., his section Bitectae 
(outer layer as above overlying a tissue of longitudinally arranged conglutinated hyphae) was 
similar to Euramalina Stizenb. He made a third division Ecorticatae (cortex composed of a layer 
of longitudinally oriented conglutinated hyphae which formed a closed ring). Ramalina arabum 
(Ach.) Meyen & Flotow is the type for Ecorticatae. Choisy (1954) commented that this thallus 
structure was similar to Alectoria Ach. 

Howe (1913-14) used spore characters to distinguish the sections he created within the genus 
Ramalina as he thought this represented a more natural classification. Section Ellipsosporae R. 
H. Howe contained species with ellipsoid or oblong spores (9-20 x 3-5-7-5 /xm) and section 
Fusisporae R. H. Howe contained species with long fusiform spores (16-35 x 3-7 /xm) and 
section Bistortae R. H. Howe contained a species with sigmoid spores. 

Each of these sections he divided into series and these series correspond in the main with 
previous divisions, e.g. Howe's series Desmazieria (Mont.) = section Corticatae Steiner = genus 
Niebla Bowler & Rundel. Series Myelopoeae = part of Myelopoea Vainio, series Fistulariae = 
stirps Fistularia Vainio = genus Fistulariella Bowler & Rundel. He made three other series, 
Tenuicorticate, Ciliate, and Fusisporae. This classification is not used. 

Du Rietz (1926) relegated the genus Desmazieria Mont, to subgenus Desmazieria (Mont.) 
(containing R. ceruchis, R. homalae, etc.) and Euramalina Stizenb. to subgenus Euramalina 
(Stizenb.), which he divided into section TenuicorticataeR. H. Howe and section fi/fecfae Steiner. 
Section Tenuicorticatae contained species without a chondroid element and this section he 
further divided into subsection Solidae Du Rietz (species with a solid thallus, containing R. 
evernioides, etc.) and subsection Tubulosae Du Rietz (species with inflated fistulose thallus: R. 
manw Mont. = Cenozo^/aMassal.). 

Du Rietz maintained section Bitectae Steiner (= Euramalina Stizenb.) but divided it into two 
subsections (a) subsection Myelopoea Vainio thallus solid, and (b) subsection Fistularia Vainio 
thallus inflated, fistulose. These extra subsections make the classification compUcated although 
the basic criteria used for division are sound. 

Zahlbruckner (1907) combined the sections of both Stizenberger (1862) and Steiner (1904) 
and made three divisions in the genus Ramalina: (i) section Ecorticatae Steiner (ii) section 
Corticatae Steiner (which incorporated Cenozosia Massal. and Desmazieria Stizenb.) and (iii) 
section Euramalina Stizenb. (= Bitectae Steiner). Zahlbruckner subdivided Euramalina 
Stizenb. using the Vainio names Fistularia Vainio and Myelopoea Vainio and series Teretius- 
culae Vainio and Compressiusculae Vainio. 

Present day taxonomists are still divided in their acceptance of which species should be placed 
in the genus Ramalina sens. str. Bowler & Rundel (1977) established the new genus Fistulariella 
Bowler & Rundel for those species originally placed in Ramalina stirps Fistularia Vainio. They 
placed emphasis on the hollow, usually perforate thallus as traits warranting genus status, yet 
Australian material belonging to stirps Fistularia Vainio present a broad range of morphology 
varying from almost totally compressed to totally fistulose. It is therefore regarded here that the 
trait of inflation of the thallus should have no greater emphasis placed upon it than the roundness 
of branches. 

Rundel & Bowler (1978) proposed a new generic name Niebla to replace the genus name 
Desmazieria Mont, as they found the former name was a homonym for the earlier described 
genus Desmazeria Dumortier (Poaceae), and 13 taxa were transferred by them from Desma- 
zieria and Ramalina into Niebla Rundel & Bowler. 

Of the various divisions put forward up to the present day, the taxonomic system used by 
Vainio would most suitably classify the Australian material, e.g. the Australian Ramahnae 
would broadly divide into three groups: (1) fistulose thaUi (2) terete and solid thalli and (3) 
compressed and solid thalh. 

(2) THE FAMILY RAMALINACEAE 

Agardh (1821: 93) erected the family Ramalinaceae ['Ramalineae'], in which he placed the genus 

Ramalina. Since that time the genus has been alternatively included with IJsnea and other 



RAMALINA IN AUSTRALIA 117 

fruticose genera in the family Usneaceae, or isolated in the family Ramalinaceae. Table 2 shows 
the various classifications which record Ramalina in RamaUnaceae (with or without other 
members of Usneaceae) and the diversity of opinions as to the position of the genus. 

Apparently Zahlbruckner (1907) and Smith (1921) did not regard septate spores nor a cortex 
with mechanical tissue sufficiently different to separate Ramalina from other genera in the 
family Usneaceae; but Watson (1929) regarded spore septation as most important and placed 
Ramalina in the Ramalinaceae. 

With modern day improvement in techniques of microscopy and chemistry together with a 
better understanding of the fungal/algal composition, the natural relationships which exist 
between the genera are being more fully understood. 

Follmann & Huneck (1969) placed three genera in the family Ramalinaceae based on their 
chemotaxonomy and anatomy (viz. Ramalina, Ramalinopsis , and Desmazieria). Poelt (1974) 
also regarded these three genera as forming a natural entity and placed them in Ramalinaceae, 
but he added a fourth genus which was not closely related, e.g. Speerschneidera. Culberson & 
Culberson (1970) pointed out that chemical evidence supported segregation of Ramalinaceae as 
a separate family as no species in the Usneaceae proper produces orcinol meta-depsides, a 
category of substances richly represented in Ramalina. 

Henssen & Jahns (1973) have kept the genus Ramalina in the family Ramalinaceae, because 
of its two-celled spores and exobasidial conidiophores, and because of the specific lichen 
substances it contains. They stated that the structure of the apothecium of Ramalina resembles 
that of the Parmeliaceae in that there is a more or less complete algal layer beneath the 
hymenium, but the Ramalinaceae differ from the characteristic cupular exciple of the 
Parmeliaceae in that the algal layer is next to, not separated from the loosely interwoven 
subhymenium. 

Keuk (1979) divided the genera in Ramalinaceae into two groups based on anatomical and 
chemical differences, and colour of pycnidial walls (a) Ramalina, Ramalinopsis, and Tricho- 
ramalina and (b) Niebla (= Desmazieria Mont, and Cenozosia Massal.). 

The most modern division within the family Ramalinaceae has been based on cortical 
diversity. Bowler (1981) defined seven genera in the RamaUnaceae: Ramalina Ach., Fistu- 
lariella Bowler & Rundel, Niebla Rundel & Bowler, Ramalinopsis (Zahlbr.) FoUm. & Huneck, 
Trichoramalina Rundel & Bowler, Dievernia M. Choisy, and Cenozosia Massal. Of these, the 
genus Ramalina contains the largest number of taxa. 

It can be seen from Table 2 that all recent authors who have specifically dealt with the question 
of relationships of genera within the two families Ramalinaceae and Usneaceae, have con- 
sidered them to be distinct morphologically, chemically, and taxonomically. 



Collectors of Australian Ramalinae 

Because the early explorers in Australian waters sailed in the temperate latitudes, it was only to 
be expected that the first Ramalina collected in Australia would be a temperate region species; 
this collection was made by Labillardiere, a botanist on the French ship 'Recherche' which sailed 
along the Great Australian Bight and visited Tasmania in 1791 (Ducker, 1979). This first 
specimen, held at H, (H-NYL 37226) was annotated 'Ramalina leiodea Nyl. Labillardiere, C. 
van Dieman'. This was an erroneous identification which has now been determined as R. inflata 
J.D. Hook. & Taylor. 

In 1802, Robert Brown in his voyages with Matthew Flinders collected R. inflata from Port 
Jackson (Sydney, N.S.W.), but this specimen was determined as Lichen fastigiatus Ach. by 
Crombie(1880). 

When J. D. Hooker accompanied Sir James Ross in his Antarctic expedition in the ships 
'Erebus' and 'Terror' in 1839, he collected the type of R. inflata J.D. Hook. & Taylor on the 
Lord Auckland Islands and from New Zealand he collected the type oiR. geniculata J. D. Hook. 
& Taylor. They also visited Tasmania and collected material which was later published in Flora 
Tasmaniae (1859). In this work Churchill Babington cites R. tasmanica Nyl. as a new species but 



118 



G. NELL STEVENS 



Table 2 The classification of genera within the two families Ramalinaceae and Usneaceae as proposed by 
various lichenologists from 1814 to the present day. 



Author 



Date Class/Order 



Tribe/Family 



Genus 



Acharius, E. 



1814 Class 

Homothalami 

Order Primus Scutellati 

Order Secundus Peltati 



Ramalina, Alectoria, Collema 
Usnea, Cornicularia 



Agardh, C. 



1821 



Ramalineae 



Ramalina 



Eschweiler, F. G. 



1824 



Usneaceae Usnea, Ramalina, Alectoria ± 

Cornicularia, Dactylina, 
Desmaziera, Endocena, Evernia, 
Everniopsis, Letharia, 
Neuropogon, Oropogon, 
Siphula, Thamnolia. 



Montagne, J. F. C. 1852 



Tr. Parmeliacea 
subtribe Usnea 



Ramalina, Usnea 



Nylander, W. 1857 Lichenacei 

Ser. Ramalodei Tr. Ramalinei 

Tr. Usneei 



Ramalina, Alectoria, Evernia, 

Dufourea, Dactylina 
Usnea, Neuropogon, Chlorea 



Stizenberger, E. 1862 Class Tr. Usneaceae 

Homothalami Ramalineae 

Usneaceae 



Ramalina, Dactylina, Cetraria, 

Evernia, Dufourea 
Usnea, Alectoria, Oropogon, 

Argopsis 



Vainio, E. A. 



1890 Order 

Cyclocarpeae 



Tr. Parmelieae 



Ramalina, Usnea, Alectoria, 
Evernia 



Hue, A. M. 



1901 Series 

Cyclocarpae 



Radiatae 
Tr. Ramlineae 
Tr. Usneae 
Tr. Alectoreae 



Ramalina, Dufourea 

Usnea 

Alectoria 



Zahlbruckner, A. 



1907 Order 

Cyclocarpeae 



Usneaceae Ramalina, Usnea, Alectoria, 

Oropogon, Dufourea, Evernia, 
Everniopsis, Endocena, 
Letharia, Thamnolia, 
Siphula, Dactylina. 



Smith, A. L. 



1921 subseries Usneaceae 

Cyclocarpineae 



Ramalina, Usnea, Alectoria, 
Evernia, Letharia, 
Everniopsis, Oropogon, 
Thamnolia, Siphula, Dactylina, 
Dufourea, Endocena 



Watson, W. 



1929 Order 

Parmeliales 



Ramalinaceae Ramalina 

Usneaceae Usnea, Alectoria, Letharia, 

Evernia, Everniopsis, Dactylina, 
Dufourea, Thamnolia, Siphula, 
Endocena 



Rasanen, V. J. P. B. 1943 Order 

Cyclocarpeae 



Usneaceae Ramalina, Usnea, Alectoria, 

Evernia, Letharia, 
Everniopsis, Dufourea, 
Dactylina, Oropogon, Ramalea 



Choisy, M. 



1957 



Ramalinaceae Ramalina 

Usneaceae Usnea, Desmazieria, Cenozosia 



RAMALINA IN AUSTRALIA 



119 



Table 2 - cont. 










Author 


Date 


Class/Order 


Tribe/Family 


Genus 


Follmann, G. & 
Huneck, S. 


1969 




Ramalinaceae 


Ramalina, Ramalinopsis, 
Desmazieria 


Henssen, A. & 
Jahns, H. M. 


1973 


Order 
Lecanorales 
suborder 
Lecanorineae 


Ramalinaceae 


Ramalina 



Poelt, J. 



1974 Order 

Lecanorales 

suborder 

Lecaniorieae 



Ramalinaceae Ramalina, Ramalinopsis, 

Desmaziera, Speerschneidera, 

Usneaceae Usnea, Alectoria, Bryopogon, 

Cornicularia, Evernia, 
Everniopsis, Himantormia, 
Letharia, Sulcaria, Neuropogon, 
Oropogon 



Bowler, P. A. 



1981 



Ramalinaceae 



Ramalina, Fistulariella, 
Trichoramalina, Niebla, 
Dievernia, Cenozosia, 
Ramalinopsis 



examination of this material during the present study has shown this specimen to be yet another 
morphotype of the very variable R. inflata. 

The Reverend Dr. W. Woolls, a school master at Parramatta, N.S.W., in 1832, collected in 
that area, and one Ramalina specimen called R. gracilis Nyl. (MEL 9432) is actually R. exiguella 
Stirton. 

During the period 1842-46, J. P. Verreaux (a French ornithologist) who was sent to Australia 
by the Museum of Natural History in Paris, collected widely in Tasmania and mainland 
Australia. In his botanical collections were several Ramalina species which he had obtained 
during that period, either by collecting them himself or by purchase of material from others 
(Lamy, in litt.). Because of this practice of obtaining specimens from other collectors there is 
uncertainty about the locality given by Nylander for two 'type' specimens he records as collected 
by Verreaux from Swan River, Western Australia, these are R. australiensis Nyl. and R. 
confirmata Nyl. Both of these species grow in eastern Australia, and the Western Australian 
types are the only material collected from that area. Nylander himself appeared unsure of the 
locality of R. australiensis as he noted: 'in Australia (prope Swan River, ni fallor) legit Verreaux 
1846'. Verreaux collections held at PC indicate he travelled north as far as Moreton Bay, 
Queensland, and collected several Ramalina species from mangroves. 

The first woman recorded as a lichen collector was Amalie Dietrich, who collected hchens in 
Queensland for a wealthy Hamburg merchant named Godeffroy, for his private museum. Some 
of these collections are now housed at Miinchen. They are collections of Ramalina species from 
the Rockhampton area. 

Friedrich Ludwig Leichhardt travelled through Queensland on his way to Port Essington 
(Darwin), 1844-45. He collected a Ramalina (now R. filicaulis N. Stevens) along the way but 
gave no exact location. As the occurrence of this species is rare today, more information about 
the locality would have been helpful in the present study. 

Ferdinand von Mueller, Victorian Government Botanist (1853-96), received lichen speci- 
mens from various collectors in Australia and these specimens were sent to overseas lichenol- 
ogists for determination, e.g. Anton Krempelhuber, Jean Miiller [Arg.], and James Stirton. 

Daniel Sullivan, a headmaster, collected lichens for von Mueller, one being the 'type' of R. 
glaucescens Krempelh., from Mt Ararat, western Victoria, and another was R. leiodea var. 
fastigiatula Miill. Arg. 'type' from the Grampians. Another Ramalina which Sullivan collected 
from Mt Ararat area (MEL 9471) now bears the name R. fimbriata Krog & Swinscow, a species 
common in East Africa. Charles French, a plant propagator at Melbourne Botanical Gardens 



120 G. NELL STEVENS 

(Filson, 1976), collected the type of/?, inflata yar.fissa Miill. Arg. from King Island, Bass Strait. 
The type of/?, lacerata Miill. Arg. was collected at Eucla, Western Australia, by J. Oliver. 

Collectors in New South Wales for the Victorian Botanical Department included T. White 
who collected the type of R. myriodada Miill. Arg. from Twofold Bay, New South Wales. 
Charles Knight, a New Zealand surgeon, collected lichens in the Sydney area; he named one 
specimen R. subgeniculata C. Knight, which was subsequently changed to R. knightiana Zahlbr. 
Knight also named R. minuscula var. alba a specimen collected by James Keys at Mt Perry, 
Queensland. This name was pubUshed by Bailey (1886), but no description was given and later 
John Shirley (1888) described it. 

Messrs. Pentzke and Hartmann were two collectors in Queensland for the Victorian Botanical 
Department (Wilson, 1889). Pentzke collected the type material of R. farinacea var. nervulosa 
Miill. Arg. from Daintree River, north Queensland, and C. H. Hartman of Toowoomba 
collected the type material of R. geniculata var. compacta Miill. Arg. 

Collections made by F. M. Bailey, Government Botanist in Queensland, John Shirley, a 
school inspector, and Mrs M. Thozet, were sent to James Stirton, W. Leighton, Charles Knight, 
or Jean Miiller [Arg.] for identification (Wilson, 1889). It was Stirton who named the two 
Ramalina types R. exiguella Stirton and R. perpusilla Stirton. Miiller Arg. described a 
Rockhampton specimen collected by Mrs Thozet as R. farinacea var. squarrosa Miill. Arg. The 
type of R. farinacea var. dendroides Miill. Arg. was collected by E. Forde from the Hunter 
River, New South Wales. Both of these taxa have now been found to be morphotypes of R. 
peruviana Ach. 

The Reverend F. R. M. Wilson, from Kew, Victoria, collected prolifically (1877-97) in 
eastern Australia, from Tasmania to Brisbane, and more particularly along Port Phillip Bay and 
Gippsland, Victoria (Filson, 1976). He collected the type of/?, calicaris var. australica Rasanen 
from Barwon Heads. Wilson himself described several taxa of Ramalina, e.g. /?. unilateralis F. 
Wilson, /?. brevis F. Wilson, and /?. brevis var. brevissima F. Wilson. 

More recent collectors of Ramalinae in AustraUa are J. H. Willis, Assistant Government 
Botanist at the National Herbarium Melbourne (Filson, 1976), A. C. Beauglehole, who 
collected Ramalinae in South AustraUa and Victoria, J. S. Whinray, who made an extensive 
survey of the lichen flora of the Bass Strait Islands, and G. C. Bratt and J. A. Cashin, whose 
collections of Tasmanian lichens have proved invaluable in the present study. 

Other present day lichenologists who have collected Ramalina specimens which have been 
used in the present study are A. Archer (N.S.W.), J. A. Elix (A.C.T.), R. Filson (Vic), G. 
Kantvilas (Tas.), R. W. Rogers (Qld.), N. Sammy (W.A.), C. Scarlett (Qld), and R. Seppelt 
(S.A.). 

Materials and methods 

The source of material for this research was threefold, (a) Personal collections (numbering over 
10,000 specimens) , the majority from Australia, with small collections from the Pacific Islands of 
Fiji, New Caledonia, Rarotonga, Tahiti, New Hebrides, New Zealand, and Hawaii, (b) 
Herbarium material from institutes, as acknowledged, (c) Herbarium and living material from 
private collections, as acknowledged. 

Most of the specimens examined during this study were tested by thin-layer chromatography 
using the techniques of Culberson (1972). Solvents B and C proved to be the most useful solvents 
to separate the acids found in the various species of Ramalina. The chemistry of type material 
was also checked. Examination of the surface features of the thallus (e.g. pseudocyphellae, 
soralia, and soredia shapes) and spore size and shape, was carried out by use of the Scanning 
Electron Microscope (SEM) using either a Cambridge or a Phillips 505. In order to observe the 
shape and thickness of the hyphae composing the outer cortex and the supportive tissue thin 
sections were examined by TEM, using an AEI Corinth 275 operated at 60 kV. Preparation of 
thallus material involved fixation of the tissue in glutaraldehyde, post fixation in osmium 
tetroxide, dehydration and then infiltration and embedding with Spurrs medium. The results 
were not always successful as the tissue tended to pull away from the medium along the outer 
cortex junction. 



RAMALINA IN AUSTRALIA 121 

All climatic data were derived from Meterological Bureau records {Climatic averages of 
Australia, 1975). 

Results 

Morphology 

(a) Soralia and soredia: Soralia are defined herein as those areas of thallus where clusters of 
soredia are found, and the term soredia is applied to the more or less spherical bodies composed 
of algae surrounded by short hyphae which act as vegetative diaspores. These vegetative 
reproduction bodies are produced by 11 of the AustraUan Ramalinae, and their constancy in 
occurrence enables this attribute to be used taxonomically. 

The position and form of the soralia are also useful criteria in the separation of each sorediate 
taxon. Round or ellipsoid soralia are produced by R. pacifica, R. caespitella, R. nervulosa var. 
nervulosa, R. nervulosa var. luciae, and R. nervulosa var. dumeticola (Plate 1 , fig. 3) ; these occur 
marginally and laminally. Ramalina peruviana and R. tenella produce small, punctiform soralia 
(Plate 1, figs 1, 2) which are mainly apical in R. tenella and marginal or lateral in R. peruviana. 
The soralia found in the other four taxa are produced in several ways, e.g. (i) by disintegration of 
the lower cortex {R. fimbriata, Plate 1, fig. 5) (ii) by the partial separation, both marginally and 
apically, of the upper and lower cortices {R. canariensis) (iii) by lack of a continuous lower 
cortex {R. reducta, Plate 1, fig. 4) and (iv) by the splitting or disintegration of parts of the lower 
cortex at intervals along the branches {R. unilateralis , Plate 1, fig. 6). The soraUa terminology 
used by Du Rietz (1924) and Beltman (1978) was not generally appHcable to the Ramalinae. 

The composition of the soredia was examined under SEM; the algal cells were not discernable 
but the fragments of hyphae which make up the outer surface of the soredium were observed to 
differ in form in several of the taxa. The northern hemisphere species R. capitata is regarded as 
having soredia covered by an 'epicortex-like' layer (James, pers. comm.) (Plate 2, fig. 1), and 
such a structure was looked for in the Australian sorediate taxa. Four species were found to 
possess some semblance of an outer covering (Plate 2, figs 2, 3, 4, 5). Whether this layer should 
be compared to the 'epicortex' which occurs in some taxa in the Parmeliaceae as described by 
Hale (1973, 1981) is debatable; but it can be likened to the 'Kittsubstanz' (cementing substance) 
mentioned by Peveling (1970). Such a layer may act as a protection against wetting of the 
soredia. 

The Australian taxa which produce soredia with this outer covering are R. caespitella, R. 
reducta, R. tenella, and R. fimbriata, and even in these only a minority of the soredia show it 
clearly. The soredia of R. caespitella are small, spherical, and have a partially smooth surface 
(Plate 2, fig. 2). Soredia of R. reducta are irregular in shape and size but fusion of the external 
hyphal segments was apparent (Plate 2, fig. 3). Ramalina tenella soredia are small but well 
defined, and some have a distinct outer covering (Plate 2, fig. 4). Ramalina fimbriata soredia are 

Plate 1 (overleaf) SEM photographs showing soralia shapes in some of the Australian Ramalinae. Fig. 1 
Apical, punctiform soralia - R. tenella. Scale 10 mm = 530 fim (x 19). Fig. 2 Marginal/lateral, 
punctiform soralia -/?./7erMViflrta. Scale 10mm = 150)u-m(x 65). Fig. 3 Round and ellipsoid soralia-/?. 
nervulosa var. dumeticola. Scale 10 mm = 220 /xm (x 44). Fig. 4 Exposed lower surface with chondroid 
strands of cortical tissue across area of soralia-/?. reducta. Scale 10 mm = 270 /lim (x 36). Fig. 5 Patches 
of exposed medulla by disintegration of the under side -/?. fimbriata. Scale 10 mm = 200 ^tm (x 51). Fig. 
6 Lower surface split apart and eroded at intervals - R. unilateralis. Scale 10 mm = 530 fim ( x 19). 

Plate 2 (overleaf) SEM photographs showing soredia forms in some of the Australian Ramalinae. Scale 
indicated by black bar. Fig. 1 R. capitata - showing smooth outer layer partially covering the soredium. 
Scale 10 mm = 22 fjun (x 440). Fig. 2 R. caespitella - showing portion of soredia covered by a smooth 
layer. Scale 10 mm = 22 /tm (x 440). Fig. 3 R. reducta - showing soredia with smooth surface (x 364). 
Fig. 4 R. tenella - showing thin layer partially enclosing the soredium (x 1236). Fig. 5 R. fimbriata - 
showing thin layer totally enclosing soredium (x 958). Fig. 6 R. unilateralis - showing soredium with a 
'woolly' appearance; no outer layer present (x 1527). Fig. 7 R. nervulosa var. dumeticola - showing 
soredia with irregular surface (x 873). Fig. 8 R. canariensis - showing soredia with a 'woolly' 
appearance; no outer layer present (x 909). 



122 



G. NELL STEVENS 







Im^b ^'^I 


lOn^B^iu^ 


^IB^'flaH 


^^^■■^S^^l 


^k^Bm 


^Hff 'fl^^^l 


^^^r^'^^i 


KJca^B^^B 


(■J^^^^^^^^^Si* 


» IbH 


■ 


^HH^^B^^H 


i^fe 


bl 


^^^^St^l 


r ^^1 


g^^B 




^^^^BIH 


Una 




Plate 1 



RAMALINA IN AUSTRALIA 



123 





^^^^^^^^^^PP^F ' .rfiT* «> i«.7/^^^^^^^H 






Bs^^^^^^^^BikL^:.!^^i^^MiH^^^^^^^^P^BI 




Plate 2 



124 G. NELL STEVENS 

large and spherical, and the formation of an external layer could be discerned on many of the 
soredia at various stages of development; the most advanced layer formation is shown in Plate 2, 
fig. 5. All of these taxa are regarded as having granular soredia. 

The soredia produced by R. unilateralis (Plate 2, fig. 6) and by R. canariensis (Plate 2, fig. 8) 
are small in size and are composed of both thick and thin pieces of hyphae; there is no external 
fusion of the hyphae so the outer surface remains loosely interwoven giving a woolly appearance 
to the soredia (Plate 2, fig. 6). The soredia are farinose. The four taxa which belong in the R. 
farinacea complex {R. pacifica, R. nervulosa var. nervulosa, R. nervulosa var. luciae, and R. 
nervulosa var. dumeticola; Plate 2, fig. 7; Plate 3, figs 1, 2, 3) also produce soredia which have an 
irregular surface made up of separate hyphal pieces which are not fused (Plate 3, fig. 2), and the 
soredia are farinose. Ramalina peruviana produces soredia which have an irregular surface 
(Plate.3,fig. 4). 

Fibrils often arise from mounds of farinose soredia (Plate 1 , fig. 2) and these features have the 
same structure as the soredia (Plate 3, fig. 7). V 

(b) Apothecia: The production of apothecia as a sexual means of reproduction occurs in all but 
four of the Australian Ramalinae (i.e. seven of the 1 1 sorediate taxa also produce apothecia , but 
only rarely, and often the mature ascospores are few and difficult to find). The esorediate taxa 
generally produce numerous apothecia. 

The position of the apothecia on the branches can be of taxonomic importance, being 
terminal, subterminal, marginal, lateral, or laminal. Terminal refers to the apothecia at the end 
of a branch. Subterminal refers to apothecia positioned near the end of the branch, with the 
branch extending beyond the apothecium, forming a short attenuate branchlet or spur, which is 
usually bent at an angle in relation to the main branch. If the spur grows long it makes the branch 
appear geniculate. Marginal refers to apothecia on the edges of compressed branches, and 
laminal refers to apothecia occurring on the surface of the thallus. If branches are subterete to 
terete the apothecia occur laterally. When laminal apothecia occur specifically at the base of 
bifurcating branches they are referred to as positioned at the axil of the branch. 

Apothecial shape varies from concave to plane to convex; in some species the apothecia 
remain concave to maturity, in others the apothecia are concave to plane when immature, but 
are markedly convex at maturity. The thalline margin is usually distinct and entire, occasionally 
becoming incised at maturity; when the disc becomes convex the margin is often obscured. 

The size of the apothecia varies greatly from 0-2 mm to 5-0 mm with exceptions to 10 mm. 
Most apothecia are shortly stalked, but sessile apothecia are found in R. australiensis and R. 
filicaulis. The colour of the disc is usually yellow-green and pruinose; however, taxa in the R. 
celastri complex have been found with orange coloured discs. 

The apothecial tissue comprises a colourless hymenium formed of asci and paraphyses. The 
asci are clavate to subcylindrical and lie amongst sparsely branched paraphyses over a developed 
hypothecium. This hypothecial layer is usually brownish-yellow as is the epithecium [as used in a 
lichenological context - Henssen & Jahns (1973)]. 

(c) Spores: The number of spores per ascus is eight, usually distichous in arrangement (Plate 5 , 
fig. 5), hyaline, 1-septate, varying considerably in size and shape, being broadly ellipsoidal to 
fusiform and either straight to slightly curved or reniform (Plates 4 & 5). 

Within a single ascus spores can vary from straight to curved, a feature also noted by Krog & 

Plate 3 SEM photographs showing soredia forms ancj, surface features in some of the Australian 
Ramalinae. Scale indicated by black bar. Fig. 1 R. nervulosa var. luciae - showing soredia with irregular 
surface (x 750). Fig. 2 R. nervulosa var. nervulosa - showing fragments of hyphae making up the outer 
surface, without a surface layer ( x 945). Fig. 3 R. pacifica - showing soredia with an irregular surface ( x 
525). Fig. 4 R. peruviana - showing soredium with an irregular surface (x 1300). Fig. 5 Punctiform 
pseudocyphellae - ^. subfraxineavar. confirmata (x 75). Fig. 6 Enlargement of section of Fig. 5 (x 750). 
Fig. 7 Fibril produced from a mound of soredia - R. peruviana. Scale 10 mm = 30 ^tm (x 341). Fig. 8 
Tuberculate pseudocyphellae - R. subfraxinea var. subfraxinea. Scale 10 mm = 30 /im (x 341). 



RAMALINA IN AUSTRALIA 



125 




126 G. NELL STEVENS 

Swinscow (1975) and Landron (1972) (Plate 4, fig. 4); therefore the shape of spores was not used 
as a taxonomic character. The size of spores has also been found to be of Hmited taxonomic value 
as most mature spores of the Ramalina species lie within the size range (8-)10-16 x 3-5-6 fxm 
(Plate 4, figs 2, 3, 4, 5, 6). The relationship between the width and length of spores, as suggested 
by Krog & Swinscow (1974) was investigated in this study to see if this could be a taxonomic tool 
for separation of closely related taxa; however exceptions could always be found in the spore 
ratios which negated the usefulness of this characteristic (Plate 5, figs 2,3,4). 

Although the spores are 2-celled, some of the spores in the size range 14-16 x 4)Ltm appear 3 
or 4-celled. These additional divisions are not produced by distoseptae so must be dense 
cytoplasmic strands (which take up the cotton blue stain) across the cell which produce a 'mock 
septation'. 

The spores of each taxa examined under SEM showed surfaces which were remarkably 
smooth (Plates 4 & 5) except for a film which seemed to coat some of the spores or remained in 
patches attached to the spore surface (Plate 4, fig. 1; Plate 5, figs 1, 2, 4, 6). 

(d) Pycnidia: The occurrence of pycnidia amongst the Australian Ramalina species is not 
common. When present, they are always pale in colour, and produce pycnoconidia which are 
rod-shaped, usually 3-5 x 0-5 /xm in size. It was observed that they mainly occurred on sterile 
thalli rather than on thalli bearing either apothecia or soredia. This suggests that they may 
provide an alternative means of propagation for the species. Their irregular occurrence and 
apparent uniformity prevented their taxonomic use. 

(e) Pseudocyphellae: Many lichenologists have discussed these morphological characters (Du 
Reitz, 1924; Culberson & Culberson, 1968; Duncan, 1970; Beltman, 1978; Bowler, 1981; and 
Hale, 1981). It is generally accepted that these features are cortical pores in the surface of the 
thallus which allow the exchange of gases. They are areas which have direct contact with the 
medullary hyphae. In the Ramalinae they vary in shape and size and several names have been 
applied to such surface features: striae, striations, tubercules, and papillae. 

Pseudocyphellae appear as shallow depressions (striae) in the cortex (Plate 3, figs 5, 6) or as 
white linear markings (striations) on the surface (Stevens, 1983fl: figs IB-D, 2A-B). In the 
Ramalina subfraxinea complex in AustraUa some of the taxa produce tubercles resembling 
raised warts, the tips of these structures containing small pseudocyphellae (Plate 3, fig. 8). The 
term 'tuberculate pseudocyphellae' was used by Krog & Swinscow (1975) to describe this form 
of pseudocyphellae. Similar surface features occur in several North American taxa, but are 
referred to as papillae, as in R. sinaloensis (Bowler & Rundel, 19726) and JR. complanata 
(Landron, 1972). 

Most sorediate Ramalina species produce pseudocyphellae of one kind or another; in the case 
of the R. farinacea taxa from the tropics, it was found that the elongate pseudocyphellae were 
precursors to soralia formation (Stevens, 1983fl). 

Bowler (1981) reported that pseudocyphellae are mainly absent in the Fistulariella taxa and 
this was found to be so in the Australian fistulose material. As a diagnostic character for the 
separation of taxa, pseudocyphellae have been found to be of little value. 

Anatomy 

The anatomical structure of all Ramalina species found in Australia comprises (1) a cortex 
(which I regard as both the outer cortex plus the inner supportive tissue) and (2) a medulla 
(which includes the phycobiont). 



Plate 4 SEM photographs showing spore shapes in the Australian Ramalinae. Fig. 1 R. subfraxinea var. 
/e/orfea- single spore. Scale 10mm = l-2/x.m(x 8000). Fig. 2^. australiensis spores. Scale 10mm = 4/Am 
(x 2400). Fig. 3 R. pacifica spores. Scale 10 mm = 4 fim (x 2400). Fig. 4 R. subfraxinea var. confirmata 
spores. Scale 10 mm = 4 /xm (x 2400). Fig. 5 R. subfraxinea var. norstictica spores. Scale 10 mm = 4 ^tm 
(x 2400). Fig. 6 R. subfraxinea var. subfraxinea spores. Scale 10 mm = 4 /xm (x 2400). 



RAMALINA IN AUSTRALIA 



127 




128 



G. NELL STEVENS 




RAMALINA IN AUSTRALIA 129 

(1) Cortex 

In the Australian taxa the outer cortical layer is usually 20 /am in thickness and is composed of 
thick walled, transverse or randomly oriented hyphae; it may be indistinct or even absent. The 
inner layer serves as supportive tissue and varies in thickness. It is composed of bundles of more 
or less pericUnally arranged thick walled hyphae which make up a continuous or interrupted 
sheath around the internal medullary hyphae. 

The detail in the outer cortex is often difficult to detect under light microscopy when yellowish 
granules are present; a problem pointed out by Bowler (1981). To obtain as much detail as 
possible, sections were examined under TEM. Both cortical layers of hyphae are highly 
gelatinized and a uniform distribution of lumina is apparent; each lumen is surrounded by a thick 
matrix of fibrillar material arranged concentrically (Plate 6, fig. 1). PeveUng (1974) commented 
that it is difficult to determine whether the different layers around hyphae are part of the proper 
cell wall or if they are extracellular additions to the wall. 

The change in orientation of the hyphae is apparent under TEM. In transverse section the 
randomly orientated outer layer hyphae appear elongate whereas the longitudinally oriented 
hyphae appear circular (Plate 6, fig. 2). Scattered between the hyphae are outhnes of crystals, 
which are more dense in the outer region and may be usnic acid crystals (Plate 6, fig. 3). 

(a) Anatomical structure as a means of separating taxa. Krog & 0sthagen (1980) divided the 
species of Ramalina found in the Canary Islands into four anatomical types based on the position 
of the supportive tissue: the farinacea type, the decipiens type, the bourgaeana type, and the 
duriaei type. The latter two types occur in species now placed in the genera Niebla and Dievernia 
and are therefore not relevant to Australian material. 

The majority of the Australian Ramalinae could be classed as belonging to the farinacea type 
(Figs 3 A, B), as they possess a thin outer layer of tissue and a continuous inner layer of 
mechanical tissue of uneven thickness. Variations can occur in the arrangement of this inner 
supportive tissue if it becomes partly discontinuous, leaving sections of outer tissue unsupported 
(Fig. 3C). However, this is not comparable with the 'decipiens type' where some of the strands 
anastomose across the medulla. None of the Australian taxa could be separated on this trait. 

Bowler (1981) pointed out that the Ramalina taxa could be divided arbitrarily into two groups 
based on the degree of specialization in the cortical tissue. One group with a narrow, indistinct 
outer cortex intergrading with the underlying supportive tissue and the other group with a 
distinctly double-structured cortex, but he commented that intergradation could be seen 
between the two forms. 

The Australian taxa can be divided in this way generally, but thalli within one taxon can often 
show differences in thickness of the outer cortical layer. Those taxa with a sparse or indistinct 
outer cortex are R. caespitella, R. celastri complex, R. pacifica, R. nervulosa, R. exiguella (Plate 
6, fig. 4), R. peruviana, R. reducta, and R. unilateralis. Those taxa with a comparatively distinct 
double-structured cortex are R. australiensis, R. canariensis, R. filicaulis, R. fimbriata, R. 
glaucescens, R. inflata complex, R. litorea, R. subfraxinea complex (Plate 6, fig. 5), and R. 
tenella. 

(b) Variation in the anatomical structure within the thallus. It was noticed when serial sectioning 
thalU that differences in anatomical structure sometimes occurred depending on the portion of 
the branch cut. Two forms of variation were found. The first form of variation is (i) variation in 
the alignment of the hyphae in the supportive tissue in basal sections compared to the hyphal 
arrangement in the mid and apical sections. Basal sections of R. celastri subsp. ovalis show a 
typical thin outer cortex merging with the longitudinally oriented hyphae in the supportive 
tissue, as found in the anatomy oiR. celastri subsp. celastri (Plate 6, fig. 7). Sections cut through 
the middle of the branch and towards the apex showed only a wide band of randomly oriented 

Plate 5 SEM photographs showing spore shapes in the Australian Ramalinae. Fig. 1 R. exiguella spores. 
Scale 10 mm = 4/u.m(x 2400). Fig. 2/?. /n/Zara subsp. inflata spores. Scale 10 mm = 4)u.m(x 2400). Fig. 3 
R. inflata subsp. perpusilla (inland taxon) spores. Scale 10 mm = 4 /xm (x 2400). Fig. 4 R. inflata subsp. 
perpusilla (coastal taxon) spores. Scale 10 mm = 4 /xm (x 2400). Fig. 5 R. celastri subsp. celastri spores 
within the ascus. Scale 10 mm = 8-3)u,m(x 1163). Fig. 6/?. //torea spores. Scale 10 mm = 4/i,m(x 2400). 



130 



G. NELL STEVENS 




Fig. 3 Anatomical structures of Australian Ramalina thalli : A . Cross section showing uneven thickness of 
supportive tissue in typical 'farinacea-type' anatomy; B. Cross section of R. filicaulis terete branch 
showing typical 'farinacea-type' anatomy; C. Cross section ofR. inflata subsp. perpusilla thallus showing 
variation from Fig. A - the supportive tissue being partly discontinuous; D. Algal clusters on the upper 
side only, as found in some thalli of the R. inflata group; E. Cross section of R. whinrayi showing thick 
outer cortex and irregular arrangement of supportive tissue which extends into the medullary region, 
with isolated rafts of supportive tissue also present. 



hyphae forming a network (Plate 6, fig. 6). This anomaly may be explained by the number of 
laminal apothecia that this taxon produces on both sides of the branch. The lack of supportive 
tissue and the random arrangement of hyphae could be due to the repeated formation of 
apothecial tissue from the middle of the branch to the apex. Brandt (1906) found that variations 
occurred within a single thallus, and illustrated this with serial sections along the branch of 
Ramalina curnowii. 

The second form of variation is (ii) variation in cortical structure between the upper side of the 
branch and the lower side. This was noticed by Brandt (1906) in the species R. populina. In 
Australian material such variation was found to occur in some specimens of R. inflata subsp. 
inflata and R. inflata subsp. perpusilla, where the inner cortex on the lower side is narrow, 
continuous and of even thickness, without an algal layer, but on the upper side it is partly 
discontinuous , of uneven thickness with algal clusters interspersed between the ridges (Fig . 3D) . 
This formation appears to be a modification of the normal Jarinacea type' structure, to enable 
optimal functioning of the phycobiont under particular environmental conditions. It is not a 
constant feature of these taxa, so cannot be used taxonomically. 

Another taxon which has an atypical and complex anatomy (Fig. 3E) is R. whinrayi. The 
cortex consists of a thick outer cortex and a thick, discontinuous sheath of supportive tissue 
made up of bundles with ridges extending across the ni,edulla or occurring as isolated rafts in the 
medulla; such intrusions produce islands of medullary tissue with algal clumps. This form of 
anatomy has been sighted in the northern hemisphere taxon R. capitata {= R. strepsilis). 

(2) Medulla 

The central portion of the thallus is occupied by medullary hyphae which are randomly oriented, 

either tightly compacted or loosely woven and arachnoid. In the fistulose taxa the hyphae may be 



RAMALINA IN AUSTRALIA 



131 












6 







Plate 6 TEM photographs showing anatomical features. External edge of outer cortex indicated by black 
arrow. Fig. 1 Lumen with surrounding fibrillar matrix. Scale 10 mm = 1-5 /itm (x 6770). Fig. 2 Section 
from outer to inner cortex - R. exiguella. Scale 10 mm = 1 fim (x 10450). Fig. 3 Usnic acid? crystals in 
outer cortex - R. peruviana. Scale 10 mm = 1-8 ju,m (x 5682). Fig. 4 Section from outer to inner cortex 
showing narrow outer cortex. Scale 10 mm = 5 /xm (x 2125). Fig. 5 Section from ouer to inner cortex 
showing broad outer cortex. Scale 10 mm = 1-2 fxm (x 5000). Fig. 6 L.S. of mid-section of branch of R. 
celastri subsp. ovalis. Scale 1 mm = 10 />tm (x 105). Fig. 7 L.S. of mid-section of branch of R. celastri 
subsp. celastri. Scale 1 mm = 19 jxm (x 53). (Photographs for Figs 6 & 7 by courtesy of A. Henssen.) 

almost totally lacking, or occur in patches appressed to the inner cortex. Ramalinafissa produces 
very little medullary hyphae, whereas in some R. inflata subsp. inflata specimens the central 
cavity is criss-crossed by arachnoid hyphae to the same degree as R. glaucescens, a species 
regarded as having a solid, compressed thallus. 



132 G. NELL STEVENS 

The sorediate species belonging to the R. farinacea group in Austraha have a thick fehy 
medulla compared to the R. farinacea taxa in the northern hemisphere which have a loosely 
compacted but continuous medullary layer. Other sorediate species, R. canariensis, R. 
fimbriata, R. reducta, and R. unilateralis, reveal large areas of exposed medulla which produce 
soralia. The most compacted medullary hyphae are found in R. whinrayi (Fig. 3E). 

Chemistry 

Usnic acid is found in the cortex of the Ramalinae in varying amounts but it is not used 
taxonomically as a diagnostic property. Atranorin occurs only rarely and in trace amounts so it is 
not used for taxonomic purposes either. It is the medullary substances produced by the 
Ramalinae - the depsides and the depsidones - which are important. 

(a) The orcinol depsides: These comprise both para- and meto-depsides. The para-depside 
divaricatic acid is particularly common and is found in ten Australian Ramalina species. It may 
occur as the sole medullary constituent or be accompanied by biogenetically closely related 
accessory acids such as stenosporic acid, nordivaricatic acid (bothpflra-depsides) or by sekikaic 
acid (a mefa-depside). 

Evernic acid, another para-depside occurs in one saxicolous species endemic to Australia, and 
lecanoric acid has been found as an accessory acid in this species. Evernic acid is rare in southern 
hemisphere Ramalinae, whereas in the northern hemisphere it is a relatively common con- 
stituent in this genus. 

The greatest number of lichen acids found in the Australian Ramalinae belong to the orcinol 
meffl-depside group. Fourteen substances have been identified in this study. The most common 
metabolite is sekikaic acid which is found in nine species. In marked contrast it is found in only a 
few northern hemisphere Ramalina species. Sekikaic acid may occur as the sole medullary 
constituent or be accompanied by a number of minor acids viz. homosekikaic, ramalinolic, 
4'-0-demethylsekikaic, 4'-0-methylnorhomosekikaic, and 4'-0-methylnorsekikaic acids. 

Cryptochlorophaeic acid is another orcinol mer^-depside which has been observed alone or 
may occur accompanied by paludosic and 4'-0-methylcryptochlorophaeic acids. Boninic acid 
too, occurs as a major acid with the minor metabolites 2-0-methylsekikaic, 2,4'-di-0- 
methylnorsekikaic, 4'-0-methylpaludosic, and 4,4'-di-0-methylcryoptochlorophaeic acids 
(Chester &Elix, 1978). 

(b) The ^orcinol depsidones: Only four P-orcinol depsidones have been observed in Austra- 
lian Ramalinae. Salazinic acid is present in four species and may be accompanied by trace 
amounts of protocetraric acid, but protocetraric acid does not occur as a major component in 
any of the species. Norstictic acid occurs in one coastal species and psoromic acid is present in 
one inland species. 

The number of depsidones found in the Australian Ramalinae is less than the number found in 
Ramalaina species in Europe or North America, where fumarprotocetraric acid, hypopro- 
tocetraric acid, stictic acid, protocetraric acid, norstictic acid, psoromic acid, and salazinic acid 
all occur. 

Salazinic acid occurs in both sorediate and fertile Ramalina species which inhabit the 
Australian coastline (viz. R. pacifica, R. tenella, and R. fissa). In R. pacifica and R. tenella trace 
amounts of protocetraric acid are sometimes present accompanying the salazinic acid. The 
occurrence of these two acids in the same thallus is probably due to incomplete enzymic 
oxidation of the side chain rhethyl group in the conversion of protocetraric acid to salazinic acid 
(Elix, inlitt.). 

Triterpenoids, derived from the mevalonic pathway occur in specimens of R. tenella, and 
terpenes have been found in several species but neither of these groups was utilized for 
taxonomic purposes. 

Chemical variation and species concept 

Since the chemistry of a particular species is usually constant it has proved a useful property in 



RAMALINA IN AUSTRALIA 133 

combination with morphological characters to define a lichen species. However, in some 
instances, morphologically identical taxa are found to contain different acids and the problem 
arises of how to treat them taxonomically. Elix (1982) pointed out that there are three common 
patterns of chemical variation (a) replacement type compounds (b) accessory type compounds 
and (c) chemosyndromic variation. 

(a) Replacement compounds: Replacement compounds occur in a number of Australian 
Ramalina taxa including R. whinrayi, R. glaucescens, the R. inflata complex, the R. farinacea 
complex, and the R. subfraxinea complex. The most common acids involved are divaricatic acid, 
sekikaic acid, and salazinic acid. 

The thalli of R. whinrayi produce either divaricatic acid ± nordivaricatic acid or the 
replacement compound sekikaic acid with minor metabolites (4'-0-demethylsekikaic acid and 
trace amounts of 4'-0-methyl norhomosekikaic acid). On the Tasmanian coastline both of these 
races occur in the one population, their morphology is indistinguishable, and one thallus was 
found to contain both divaricatic and sekikaic acids. 

On the Bass Strait islands only the divaricatic acid race occurs on the islands in the Hogans 
Group (the most northerly islands) , whereas on the Kents Group of islands and on Craggy Island 
only the sekikaic acid race was found, although most thalli contain trace amounts of divaricatic 
acid as well. Because of this dual occurrence of the acids in some thalli, both acid races are 
regarded as belonging to the one taxon, viz. R. whinrayi. 

An intensive study of the habitats in which each acid race occurs would be necessary to 
understand and interpret the segregation of the two acid races on the different islands. A survey 
similar to that carried out by Culberson (1969fl) on taxa in the R. siliquosa complex may solve 
this problem if it could be found that the Tasmanian population occupied separate niches 
according to the acids present. In this case the chemical races could be regarded as taxonomically 
significant (Culberson, 1967, 1969^) and the explanation of the mixed acids would infer 
hybridization (Brodo , 1978) . But the islands are small and inaccessible and such a survey was not 
possible during this study. 

The occurrence of replacement compounds was also observed in the R. inflata group, where 
lichen populations which are morphologically indistinguishable may contain either divaricatic 
acid or sekikaic acid as medullary substances. Here again such taxa have been regarded as 
chemical strains within the species; a decision strengthened by the presence of trace amounts of 
sekikaic acid in some thalli which contain divaricatic acid. A third replacement compound, 
salazinic acid, also occurs in the R. inflata complex. Biogenetically this depsidone is quite remote 
from the two depsides, yet thalli possessing this acid are remarkably similar to those with a 
depside chemistry. However, the depsidone taxon has been retained as a separate species {R. 
fissa), as it has a geographically different distribution pattern from the depside containing 
taxa. 

(b) Accessory compounds: Accessory type compounds are also present in the R. inflata 
complex, occurring sporadically in addition to the constant constituents, but having no 
correlation with any morphological or distributional variations (Elix, 1982). The presence of 
nordivaricatic acid in some divaricatic acid taxa and the presence of consalazinic or scabrosin 
derivatives in the salazinic taxon have been regarded as instances of accessory acids accompany- 
ing the major constituent and thus requiring no taxonomic recognition. Similarly the accessory 
acid connorstictic has been found in the norstictic acid taxon in the R. subfraxinea complex. 

Another Ramalina species complex comprising taxa with chemical variants of the replace- 
ment type and with accessory compounds is the tropical/subtropical R. farinacea complex. The 
acids involved are divaricatic acid (-1- accessory compound stenosporic acid) and the sekikaic 
aggregate of acids and salazinic acid, but this group is further complicated by chemosyndromic 
variation occurring within taxa which produce the sekikaic aggregate of acids. 

(c) Chemosyndromic variation: A chemosyndrome is a group of biogenetically related metab- 
oUtes where one or two compounds are regularly the major components, and the minor 



134 G. NELL STEVENS 

biosequentially related constituent of one taxon becomes the major constituent of the other 
(Culberson & Culberson, 1977; Elix, 1982). 

In the R. farinacea complex, the two tropical/subtropical taxa R. nervulosa var. luciae and R. 
nervulosa var. dumeticola contain the same acid components but these occur in different 
quantities in each taxon (Stevens, 1983a). 

Correlation between secondary-product chemistry and ecogeography in the Ramalina 
subfraxinea complex 

The importance of chemistry in identifying taxa which have different amplitudes of ecological 
tolerance has been stressed by Culberson (1967, 1969/>), Culberson & Culberson (1967), and 
Sheard (1978). The notable example of correlation between secondary-product chemistry and 
ecogeography is found in the R. siliquosa species complex of the northern hemisphere. This 
complex contains six sibling species each with a different depsidone chemistry (norstictic acid, 
salazinic acid, stictic acid, protocetraric acid, hypoprotocetraric acid, and acid deficient). 
Culberson (1969^) maintained that each chemical race was a different species which reflected 
the physiological differences between the taxa. However, the R. siliquosa chemistry comprises a 
replacement series of medullary depsidones which are able to be ranked by increasing numbers 
of oxidation steps in their biosynthesis (Culberson, Culberson & Johnson, 1977). 

The maritime R. subfraxinea complex in the southern hemisphere is similar to the maritime R. 
siliquosa complex in the northern hemisphere as it is made up of taxa which produce six different 
acids; four of the acids are depsides and two are depsidones (viz. sekikaic acid, divaricatic acid, 
cryptochlorophaeic acid, boninic acid, norstictic, and salazinic acids), but these acids are not 
closely related biosequentially (Elix, in litt.). 

Morphologically only the salazinic acid taxon is distinguishable; the other taxa show only 
slight morphological variations from one another, none of which is sufficiently unique and 
consistent that the chemical constituents of every individual can be infallibly predicted from 
appearance alone. 

Culberson, Culberson & Johnson (1977) found a correlation between the chemistry in the 
R. siliquosa taxa and different amplitudes of ecological tolerance of each taxon. Accordingly, 
a study of the thalli belonging to the R. subfraxinea complex was undertaken in order to find 
if a particular chemotype could be related to a particular type of habitat based on the 
amount of exposure to the sea (e.g. exposed = on trees or rocks facing the open sea; sheltered 
= on trees growing on the landward fringe of a mangrove community or on trees in clay pan 
habitats). 

In Table 3 the type of habitat (exposed or sheltered) at each location site is compared with the 
percentage of thalli containing each acid found at the site. The table shows that there is a 
difference in the percentage of certain acids at each site. The cryptochlorophaeic acid taxon 
occurs in areas experiencing the greatest amount of exposure to the open sea, the boninic acid 
taxon is generally found in exposed habitats (usually adjacent to quiet water), whereas the thalli 
containing sekikaic acid, divaricatic acid, or norstictic acid are mainly found in sheltered well 
protected areas away from salt spray. Salazinic acid containing thalli appear to tolerate a broad 
spectrum of habitats from exposed to sheltered. 

From Table 3 it can be seen that there is a definite tendency for different chemotypes to 
occupy some microhabitats more commonly than others. 

This environmental selection of different chemotypes into separate habitats is also apparent 
to a larger extent along the more tropical coastline of Australia (e.g. north of latitude 16°30'S), 
where collections made from any one site always contain only one chemotype. When the type of 
habitat was considered it was found that collections from Princess Charlotte Bay (a vast area of 
clay pan) contained sekikaic acid, whereas thalli collected from the exposed shorelines of the 
small tropical islands of Ingram I., Turtle I., and Sue I. all contained cryptochlorophaeic acid; 
thaUi collected from Lizard I. all contained salazinic acid. 

Although the collections from these northern regions may not have been as numerous as those 
to the south, enough material was collected at each site to make these findings plausible. 



RAMALINA IN AUSTRALIA 



135 



Table 3 Medullary metabolites expressed as a percentage of total number of specimens collected at 
various sites in relation to type of habitat and latitude. 







ta 














i 










o 










X 


1 


C3 

o 

1 

2, 


o 

a 
>^ 

H 


o 

'H 
'c 
o 
CQ 


o 

O 

u 


0) 

CO 


o 


o 

'S 

00 


CO- 

> 

5 




16 


Daintree River 


sheltered 


— 


— 


23 


— 


37 


40 


87 


16 


Mossman 


sheltered 


— 


29 


63 


— 


8 


— 


38 


17 


Mission Beach 


exposed 


— 


75 


10 


— 


15 


— 


20 


18 


Murray River 


exposed 


— 


40 


— 


33 


27 


— 


15 


18 


Hinchbrook I. 


sheltered 


1 


42 


24 


15 


16 


2 


82 


19 


Townsville 


exposed 


— 


59 


29 


12 


— 


— 


17 


20 


Bowen 


exposed 


3 


94 


— 


3 


— 


— 


33 


21 


Hallidays Bay A. 


sheltered 


— 


— 


32 


68 


— 


— 


34 


21 


HallidaysBayB. 


exposed 


— 


94 


— 


6 


— 


— 


35 


21 


Eimeo 


sheltered 


25 


35 


4 


36 


— 


— 


28 


22 


St Lawrence 


sheltered 


9 


8 


14 


69 


— 


— 


70 


23 


Keppel Sands 


exposed 


— 


96 


— 


4 


— 


— 


46 


23 


Gladstone 


sheltered 


3 


42 


30 


25 


— 


— 


60 


24 


Turkey 


sheltered 


4 


48 


48 


— 


— 


— 


33 


25 


Burnett Hds 


exposed 


10 


90 


— 


— 


— 


— 


23 


26 


Hervey Bay 


exposed 


36 


64 


— 


— 


— 


— 


22 


27 
28 


Moreton Bay 


exposed 


1 


99 


— 


— 


— 


— 


100 


29 


Clarence R. 


exposed 


66 


34 


— 


— 


— 


— 


23 



Distribution patterns in relation to climatic indices 

The distinct distribution patterns of the RamaUnae indicate that there are differences in the 
ecological requirements of each taxon. The composite map of the distribution of all of the taxa 
(Fig. 1) shows that geographically the distributions lie in either the tropical or the temperate 
zones with some taxa overlapping in the intermediate zones (e.g. subtropical and warm 
temperate). In the higher plants it has been recognized that plants growing under tropical 
conditions have different growth responses compared to plants growing under temperate 
conditions, and that such responses are closely related to thermal optima for photosynthesis 
(Specht, 1981c; Nix, 1982). 

As a knowledge of the major climatic indices which regulate plant response (e.g. temperature , 
precipitation, and solar radiation) has proved valuable in understanding the distribution of the 
phanerogram flora of Australia, it seemed reasonable to examine their value in understanding 
the distribution of the Ramalina taxa throughout AustraUa. 

Nix (1982) proposed that Australian plants can be classified in terms of their photosynthetic 
response to temperature. His three major groups are relevant to the pattern of RamaUnae 
distribution. (1) Megatherm Group - plants with photosynthetic optimum 26-28°C; lower 
threshold 10°C, upper threshold 38°C. (2) Mesotherm Group - plants with photosynthetic 
optimum 19-22°C; lower threshold 5°C, upper threshold 33°C. (3) Microtherm Groups - plants 
with photosynthetic optimum 10-14°C; lower threshold 0°C, upper threshold 25°C. These three 
major groupings have geographical equivalents as shown in Table 4. 

Nix (1982) integrated the climatic factors, temperature, precipitation, and solar radiation into 
a single multif actor Growth Index (G.I.). The Growth Index is different for plants with different 
thermal growth responses. Figs 4A-B and 5 A indicate the distribution of megatherm, 
mesotherm, and microtherm plants respectively with G.I. values >0-45 and G.I. values >0-30 



136 G. NELL STEVENS 

Table 4 Thermal divisions of Australian plant biota and their geographical equivalents, as suggested by 

Nix (1982) . 

Mean annual Geographical 
Thermal divisions temperature equivalent 



Megatherm 


24°C 


Megatherm/Mesotherm 




mterzone 


20-24°C 


Mesotherm 


14-20°C 


Mesotherm/Microtherm 




interzone 


12-14°C 


Microtherm 


<12°C 



tropical 

subtropical 

warm temperate 
cool temperate 



<0-45 in the first two groups and with G.I. values <0-6 to >0-2 in the latter. The multifactor 
Growth Index (G.I.) has values ranging from zero to unity and can never exceed the value of the 
single most limiting factor. It is apparent from Fig. 4C, that Ramalina taxa occur only in those 
areas with a G.I. larger than 0-30 in at least one thermal response category. 

The distribution pattern of each Ramalina taxon conforms in a general sense to the patterns of 
G.I. for megathermic, mesothermic, and microthermic phanerogams, indicating that the 
climatic indices which determine distribution limits for the higher plant species also create 
similar limits for the species of Ramalina. 

Biogeographical division of the Australian Ramalinae 

By superimposing the phanerogam thermal growth response patterns onto the distribution 
pattern of each Ramalina taxon, the latter are able to be divided into megatherm, mesotherm, 
and microtherm groups quite satisfactorily (Figs 4C, 5B). 

(1) The megatherm element 

Nine of the Ramalinae can be regarded as confined to a region with a megatherm Growth Index 
value >0-30 <0-45. Their southern Umits rarely extend below the tropics (latitudes 23°S-24°S). 
These taxa are R. tropica, R. tenella, R. subfraxinea var. subfraxinea, R. subfraxinea var. 
norstictica, R. nervulosa var. nervulosa, R. nervulosa var. luciae, and R. litorea. Ramalina 
subfraxinea var. confirmata and R. subfraxinea var. leiodea are exceptions as their southern limit 
reaches latitudes 33°S and 29°S respectively (Table 5). The occurrence of R. subfraxinea var. 
confirmata on Sue Island (latitude 9°20'S) confirms its placement in the megatherm group; R. 
subfraxinea var. leiodea extends to 18°S (Figs 4A, 28). 

(2) The mesotherm element 

Other Ramalina taxa which have an extended range along the eastern coastline are regarded as 
mesotherm elements as they have a southern distribution reaching the warm temperate region of 
Australia. These taxa are R. exiguella, R. inflata subsp. perpusilla, R. peruviana, and R. pacifica 
(Table 5). Although R. pacifica extends into the tropics to latitude 17°S it has been regarded as a 
mesotherm element because of its world distribution pattern which indicates its overall range is 
in the subtropics (Stevens, 1983ft). 

The presence of R. exiguella, R. peruviana, and R. inflata subsp. perpusilla in the tropical 
regions at higher elevations than sea-level is similar to the occurrence of subtropical or even 
temperate phanerogams which are reported to find an optimal thermal environment with 
increase in elevation at low latitudes (Nix, 1982). Two taxa which have restricted distribution in 
the mountains at altitudes exceeding 900 m are R. filicaulis and R. nervulosa var. dumeticola. 
Being rare in occurrence it is difficult to establish in what thermal group they belong. They have 
been tentatively placed as mesothermic plants as the former occurs in areas where R. celastri 
subsp. celastri, R. peruviana, and R. inflata subsp. perpusilla (all mesothermic plants) grow; and 
the latter must be a mesothermic offshoot of the otherwise megathermic R. nervulosa group. 

The distribution pattern of R. inflata subsp. perpusilla covers a region corresponding to 



RAMALINA IN AUSTRALIA 



137 




DISTRIBUTION PATTERN OF MEGATHERM PHANEROGAMS 
^S Growth indw n\um > 0.46 
t=l Growth indax vsluM > 0.30 < 0.46 
(Aftw Nix, 1981) 



B 



DISTRIBUTION PATTERN OF MESOTHERM PHANEROGAMS 
IMI Growth indu nluM > 0.45 
llllll Growth indn nlun > 0.30 < 0.46 
(Aftw Nix, 1981) 





Fig. 4 A. Distribution patterns of megatherm 
phanerogams with G.I. values >0-30; B. Dis- 
tribution patterns of mesotherm phanerogams 
with G.I. values >0-30; C. Ramalina distribu- 
tion pattern with megatherm and mesotherm 
plant boundaries for plants with G.I. values 
>0-30, superimposed. 



138 



G. NELL STEVENS 





5 




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■* 




<N 




■* 




T— ( 




■^ 




o 




"* 




On 




CO 




CX3 




<^ 




I^ 




r<^ 




so 




m 




m 




m 




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. tropica 

. tenella 

. subfraxinea \ar. subfraxinea 

. subfraxinea var. norstictica 

. nervulosa var. nervulosa 

. subfraxinea var. confirmata 

. litorea 


. nervulosa var. luciae 

. exiguella 

. inflata subsp. perpusilla 

. peruviana 

. pacifica 

. subfraxinea var. leiodea 


. filicaulis 

. celastri subsp. celastri 

. australiensis 


. nervulosa var. dumeticola 
. inflata subsp. inflata 
. glaucescens 


. reducta 

. inflata subsp. australis 

. fissa 

. canariensis 

. celastri subsp. ovalis 

. fimbriata 

. unilateralis 


-2 
■f ^ 



Q^D:;Q:;Q;Q;c<Qcc<c<Q:;D:;D:;Q:;c<Q:;Q:;aiQ:;Q:;Q::QCQ:;Q:;Q:;ci^ 



RAMALINA IN AUSTRALIA 139 

mesotherm response patterns with G. I. values above 0-30. The inland areas with G.I. values 
>0-30 <0-45, correspond to areas where the sekikaic acid race as well as the divaricatic acid race 
occur. Along the coast, the areas with G.I. >0-45 mainly produce thalli with divaricatic acid ± a 
trace of sekikaic acid. 

To the south, the mesotherm region with G.I. values >0-45 coincides with the R. inflata subsp. 
inflata distribution (divaricatic acid alone or rarely + trace sekikaic acid). No specimens of the 
R. inflata complex occur in the region delimited by the mesotherm elements with G.I. >0-30 
<0.45 in inland New South Wales. In the temperate southern and south-western regions 
of Australia R. inflata subsp. australis occupies this mesotherm region (G.I. >0-30 <0-45) 
and again the sekikaic acid race is present at inland sites, as well as the divaricatic acid race.. 
Along the coastline only the divaricatic acid race is present which is a similar situation to that 
found with the taxon R. inflata subsp. perpusilla. Thus in the R. inflata complex, only taxa 
containing divaricatic acid ± trace sekikaic acid are found along the eastern and southern 
coastUnes. 

In the south-west corner of Western Australia both divaricatic acid and sekikaic acid taxa 
occur, reflecting the general aridity of this part of Australia even close to the coast. 

There is a continuation of both mesotherm elements into Tasmania (Fig. 4B) and both R. 
inflata subsp. inflata and R. inflata subsp. australis occur, but in different regions. The R. inflata 
subsp. inflata taxa occur in the inland montane region whereas R. inflata subsp. australis taxa are 
found in the drier eastern areas (Figs 21, 22). Sekikaic acid taxa do not occur in Tasmania 
(extreme dryness is not a feature of the island). A microtherm influence could be responsible for 
the distribution pattern of the R. inflata subspecies in Tasmania and in many instances the 
presence of intermediate morphs causes difficulty in delimiting Tasmanian taxa. 

Another taxon placed in the mesothermic group is R. celastri subsp. celastri. The majority of 
collection sites for this taxon lie within the belt with G.I. values >0-45, with a few collection 
localities to the west in the area covered by the mesotherm belt with G.I. >0-30 <0-45. 
Specimens from the latter area are always broader than coastal specimens. This taxon extends 
south to latitude 38°S with lessening frequency. At approximately 38°S, 145°E the mesotherm 
region of plants with G.I. >0-45 is replaced by the region of plants with G.I. >0-30 <0-45 (Fig. 
4B). This latter region is occupied by R. celastri subsp. ovalis, a taxon which extends to Western 
AustraUa (Fig. 20). It is also found in northern Tasmania. 

Ramalina australiensis occurs coastally in the mesotherm pattern with G.I. value >0-45 (e.g. 
Toogoom in Queensland and Yamba to Twofold Bay in New South Wales, latitudes 25°S and 
29°-37°S respectively). 

The main distribution of R. canariensis is along the southern coastline of Australia in the 
region with mesotherm G.I. values >0'30 <0'45 which corresponds to a Mediterranean cUmate 
region. Such environments are found in South Africa, Chile, and the Mediterranean (Raven, 
1973) - all areas where this taxon occurs. The presence of/?, canariensis at several sites along the 
south-east coastUne of Australia may be due to suitable microhabitat conditions. 

The two species found on the Bass Strait islands (viz. R. whinrayi and R. caespitella) cannot be 
ascribed accurately to any of the three response groups. 

(3) The microtherm element 

There are four taxa which occur in the mountainous areas of eastern and/or southeastern 
Australia at altitudes exceeding 600 m which can be regarded as belonging to the microtherm 
response group: (1) R. reducta, (2) R. glaucescens, (3) R. unilateralis , and (4) R. fimbriata. 

(1) Ramalina reducta occurs at elevations over 1000 m in open-forests on the New England 
tableland. The area experiences very low winter temperatures with snow. Figs 5A-B show that it 
covers a region with G.I. values >0-2 <0-6. 

(2) Ramalina glaucescens is classed as a microtherm element with its two chemical races 
occupying two different response groups (i) G.I. >0-4 <0-6 and (ii) G.I. >0-2 <0-4 the former 
region being occupied by the sekikaic acid race and the latter by the acid-deficient race (Figs 5B, 
13). 

(3) Ramalina unilateralis occurs inland and along the coast at several sites, but its distribution 



140 



G. NELL STEVENS 




Microtherm 

■■GI > 0.6 

GI > 0.4 <0.6 
(SSSa GI > 0.2<0.4 

GI I 0.2 



Fig. 5 A. Distribution patterns of microtherm phanerogams with G.I. values >0-2 (after Nix, 1981). 

pattern corresponds with the microtherm element rather than the mesotherm element. Its range 
covers a region where microthermic G.I. values are >0-2 <0-6 (Fig. 5B). 

(4) Ramalinafimbriata grows on rock outcrops at high altitudes in the open-forest country of 
temperate southeast Australia (Fig. 5B); its range covers a region where microthermic G.I. 
values are >0-2 <0-6. 

Biogeography 

Taxa in the genus Ramalina found in Europe and North America are generally distinct from the 
taxa which occur in Africa, South America, Australasia, and the Pacific Islands. 

The records of Ramalina taxa which occur in the various regions of the world are scattered 
through the literature. Only the few present day revisions of the genus can be relied on to contain 
correct identification of the species; many publications record names of doubtful application. 
Those regions covered in recent reliable publications are: Canary Islands (Krog & 0sthagen, 
1980), East Africa (Krog & Swinscow, 1974, 1975, 1976), Fennoscandia and the British Isles 
(Krog & James, 1977), Europe (Poelt, 1969), and the West Indies (Landron, 1972). 

Although the Ramalinae of North America have not been completely revised, many of the 
taxa have been recently investigated by Hale (1978), Rundel (1978fl), Bowler (1977), Bowler & 
Rundel (1972a, 19726, 1973, 1974, 1978), Rundel & Bowler (1974, 1976), Hale & Culberson 
(1970), and Moore (1968). Less recent studies cover the Hawaiian lichens (Magnusson & 
Zahlbruckner, 1945; Magnusson, 1956); the Chinese Ramalinae (Zahlbruckner, 1930) and the 
Japanese RamaUnae (Asahina, 1938, 1939). 

The South American Ramalinae are in need of revision but records of taxa collected in various 
regions of that continent appear in publications of Vainio (1890), Malme (1934), FoUmann 



RAMALINA IN AUSTRALIA 



141 




Fig. 5 B. Four7?ama/mfltaxawithdistributionpatternswithinthemicrothermboundaries:l = i?. ret^Mcffl, 
2 = R. glaucescens (sekikaic acid race), 3 = R. unilateralis , 4 = R. fimbriata. 



(1967), Rundel (1978^), Osorio (1970fl, 19706, 1972, 1978), and Osorio, Aquila & Zanette 
(1980), as well as others. Papers with Ramalina taxa found on the Pacific Islands are listed in 
Appendix A of Lichen Ecology (Hawksworth, 1977). The 'Catalogue of lichens from India, 
Nepal, Pakistan and Ceylon' (Awasthi, 1965) was consulted to obtain species numbers in India, 
as no information concerning the present revision of Indian Ramalinae was obtainable from 
India. Nine Ramalina taxa are described for New Zealand (Galloway, 1985). 

From the above mentioned records and personal knowledge, some idea of the global 
distribution of the taxa which occur in Australia was obtained (Table 6, Fig. 6). Table 6 shows 
that Africa has the greatest number of species in common with Australia (50%). This close 
relationship with African lichens was mentioned by Rogers & Stevens (1981). New Zealand and 
South America have the next highest percentages of species in common with Australia. It is 



142 



G. NELL STEVENS 




Fig. 6 Global distribution of Ramalina species found in Australia. 



interesting that (as far as could be ascertained) there is little similarity of species between India 
and Australia (see below). The lack of correlation between Australian taxa and those of North 
America, Europe, and Asia, confirms that the greatest similarities are found in countries which 
have a former Gondwanaland origin. 

Palaeobiogeography 

Before presenting an interpretation of the likely palaeobiogeography of the Ramalinae, an 
understanding of some aspects of the Earth's geological history has to be appreciated. 

(1) Geological history of Gondwanaland 

The geological events that took place during the Cretaceous and Tertiary Periods which are so 
vital to the understanding of the distribution patterns of plants are summarised in Table 7. 

The early ideas of land bridges between continents and the later theory of Continental Drift 
have been superseded by the theory of Plate Tectonics. This theory is now generally accepted by 
most geologists to account for the separation of continents which show evidence of having been 
once grouped together as a Pangaea or super-continent, or as two large continents, Laurasia 
(north) and Gondwanaland (south). In this theory, the continents break up by rifting and 
separate by sea-floor spreading. New oceanic crust is formed at spreading ridges and consumed 
in subduction zones. The theory of an expanding earth (Carey, 1976) obviates the necessity for 
subduction zones and produces a more neatly-fitting Pangaea than other models, especially in 
the closer fit of northern and north-west Australia with Asia, making the distribution of now 
distant but similar biota more easily understood; unfortunately this theory has not had general 
acceptance. 

(2) Plant distribution related to geological events 

The map of the Gondwanaland continent during the Cretaceous Period (130-100 m.y. B.P.) 
[Fig. 7] shows the relevant palaeolatitudes at that time (Powell, Johnson & Veevers, 1981). 
Raven & Axelrod (1974) stated that direct migration from South America and Africa via 



RAMALINA IN AUSTRALIA 143 

Table 6 The global distribution of the Ramalina taxa occurring in Australia. 















u 


c 
























« 


2 
























'u 


"5 


a 


















•a 


c 


Oh 
(U 




Si 

c 


















c 


a 


^ 


.£: 


U 






W3 






3 




6 

< 

"3 


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0) 


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on 


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1 8 


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J2 o 


-a 

c 

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a 
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3 


s 
< 

s: 

t; 

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to 


c 
2 


Ramalina species 


< 


< 


00 


z 


1— 1 


►iSO 


^O 


H 


w 


z 


^ 


U 


R. australiensis 


* 






* 


















R. caespitella 


* 
























R. canariensis 


* 


* 


* 












« 








R. exiguella 


* 


* 




* 


















R. filicaulis 


* 
























R. fimbriata 


* 


* 




« 


















R. glaucescens 


* 






* 


















R. litorea 


* 












* 












R. peruviana 


* 


* 


* 


* 


* 


* 




* 




* 


* 




R. reducta 


* 


* 






















R. tenella 


* 


* 


* 






* 














R. unilateralis 


* 


* 


* 




















R. whinrayi 


* 
























R. celastri 


























subsp. celastri 


* 


* 


* 


* 


* 


* 




« 




* 


* 




subsp. ovalis 


* 


!|< 


* 


* 








* 










R. inflata 


























subsp. inflata 


* 


* 


* 


* 




* 








* 


* 


* 


subsp. perpusilla 


* 
























subsp. australis 


* 
























R. fissa 


* 
























R. nervulosa 


























var. nervulosa 


* 


!|C 










* 












var. luciae 


* 


* 






* 


* 














var. dumeticola 


* 


* 






* 


* 














R. pacifica 


* 


* 




* 




* 














R. subfraxinea 


























var. subfraxinea 


* 




* 




















var. leiodea 


* 










* 














var. confirmata 


* 










* 


* 












var. norstictica 


* 
























R. tropica 


* 
























TOTALS 


28 


14 


8 


9 


4 


9 


3 


3 


1 


3 


3 


1 



Antarctica to Australia was possible at this time (Table 7). Even in the late Cretaceous, 
AustraUa still had connections with South America through Antarctica, but the migration of 
plants via the southern route probably became progressively reduced until finally stopped by the 
break of Australia from Antarctica in the Eocene (53-38 m.y. B.P.). The early break-away of 
India from the rest of Gondwanaland (125-100 m.y. B.P.), and its rafting north across the 
equator with the loss of some Gondwanaland species, could account for the lack of taxa similar 
to the Australian RamaUnae. 

Two major pathways of dispersal of the higher plants from west Gondwanaland have been 
postulated by plant biogeographers: (1) a temperate migration route via Antarctica from west 
Gondwanaland (i.e. South America and Africa), to Australia and New Zealand, and (2) a 
tropical route from East Africa to Madagascar, the islands of the Indian Ocean and Indo- 
Malaysia to Australia (Croizat, 1952; Raven & Axelrod, 1974). As the Ramalina species found 
in Australia can be placed in a range of temperature response groups from cool temperate to 



144 



G. NELL STEVENS 



Table 7 The history of the break-up of Gondwanaland, indicating available migration routes (from 
Raven & Axelrod, 1974 and Powell, Johnson & Veevers, 1981). 



Million 
years BP 



Geological periods 



Event/geographic situation 



125 

110 + 10 
100 



early Cretaceous 



mid Cretaceous 



90 



80 



late Cretaceous 



63 


early Palaeocene 


55 


Palaeocene-Eocene 


49 


mid Eocene 


45 




30 


mid Oligocene 


20 


early Miocene 


10 


late Miocene 


Present 


Holocene 



Africa and South America begin to split; India begins to split 

from Australia/ Antarctica but remains attached to 

Madagascar. 

Direct migration between west Gondwanaland and Australia 

ceases. 

Direct migration between Africa and Madagascar still 

possible. 

India rafts northwards and is completely separated from 

Australia and Antarctica; no exchange of species possible. 

Migration of species between Africa and Australia via east 

Antarctica still possible. 

Africa and NE. Brazil separated by a narrow strait; migration 

possible at this point. 

Africa and South America separated by 800 km of ocean but 

numerous islands linked them along the mid- Atlantic Ridge. 

Australia ceases to have any migration from Africa. 

Australia/ Antarctica extend from 40°S to the polar region. 

South America closely connected with Antarctica 80-30 

milHon years BP. 

Separation of New Zealand and New Caledonia from 

Australia. 

Africa and Europe connected at Spain and possibly with Asia 

at Arabia, allowing both northward and southward 

migration. 

Oceanic crusts form between Antarctica and Australia; 

northern edge of Australia now at 30°S and commencement 

of separation of Australia from Antarctica. 

Separation of the continental margins of Australia and 

Antarctica but another 10 m.y. of migration of species from 

South America through Antarctica to Tasmania along the 

South Tasman Rise before final separation. 

Direct migration between India and Asia. 

The leading edge of Australia as it rafts northwards still south 

of the latitude of the Sunda Arcs. 

Initial contact between Sunda Arc and New Guinea 

(abundant precipitation). 

Collision of Australia with the Sunda Arcs. 

Antarctic ice-sheet rapid expansion (lowered precipitation). 

Australia moves towards the equator at 66 mm year~^ 



tropical, it could be hypothesised that their routes of entry to Australia were from both the south 
(the microtherm and mesotherm taxa) and from the north (the megatherm taxa). 

(3) Dispersal and migration 

Any postulates concerning the migration pattern of the Ramalinae from a centre of origin are 
purely hypothetical as no fossil records are available and any assumptions made are based on the 
migration pathways of the higher plants deduced from fossil records. The approach used in this 
study is similar to that of J0rgensen (1983) and Sipman (1983) although the writer does not agree 
with all of their conclusions. 

Biogeographic evidence suggests that many genera and families of gymnosperms and 
angiosperms had evolved before the breakup of Gondwanaland began in the Cretaceous, and 
had arrived in Australia from South America and Africa via Antarctica during the middle 



RAMALINA IN AUSTRALIA 



145 




Fig. 7 Position of east and west Gondwanaland in the early to mid-Cretaceous (130-100 m.y. B.P.). Ticks 
show longitudes which have no fixed origin (Adapted from Powell, Johnson & Veevers, 1981). 



Cretaceous (108-1(X) m.y. B.P.). Fossil evidence indicates that many of the Gondwanan 
families and genera had occupied Australia by the early Tertiary (63-55 m.y. B.P.) (Specht, 
1981c). 

(a) Step by step dispersal: Schuster (1979) stated that migration of land plants normally tends to 
be by short range dissemination as members of structured communities, and he proposed that 
the dispersal of Hepaticae occurred as part of the dispersal of whole plant communities. It is also 
reasonable to hypothesise that the Ramalina species also made this step by step spread across the 
continents of Laurasia and Gondwanaland with the migrating structural communities during the 
Cretaceous-early Tertiary. 

This suggested step by step migration of plant communities appears to be a valid means of 
dispersal as this enables plants which have similar habitat requirements to expand as a whole, in 
favourable cUmates, and so migrate when conditions are favourable and disappear from areas 
where environmental conditions have deteriorated. It seems highly likely that under the 
circumstances lichens which grow within these communities would migrate at the same rate and 
in the same direction. 

If step by step migration halted with sea-floor spreading of the continents, then another means 
of dispersal has to be proposed. Seeds of the higher plants have been found to be carried by birds 
and animals over long distances; the lighter seeds being wind dispersed. The Ramalinae produce 
both spores and vegetative diaspores and it is feasible that these could be wind dispersed or 
carried by birds. 

(b) Wind dispersal: If long distance dispersal of spores by the wind is accepted then this means 
of dispersion would have been in operation in early times. It is reported by Raven & Axelrod 
(1974) that with the separation of Australia and Antarctica in the Eocene, there was a 



146 G. NELL STEVENS 

Strengthening of the circum-Antarctic wind which could have carried spores from South 
America to AustraUa, Tasmania, and New Zealand. 

The success of a spore in a new environment would need the availability of a suitable 
phycobiont and the lack of competition from plants already in the area. With the vegetative 
diaspore both partners arrive together so that provided the environment is hospitable to their 
genetic requirements they will readily become established in the new habitat. Spores can 
become established after long distance wind dispersal since fertile lichen species have been 
found on the islands of Hawaii. J0rgensen (1979, 1983) remarked on the efficiency of wind 
dispersal of lichen spores in the population of the remote island of Tristan da Cunha. 

If the original dispersal of the RamaUnae is comparable with the original dispersal of 
angiosperms and gymnosperms, then their migration from west Gondwanaland may have taken 
place by both step by step migration and wind dispersal, with the latter means continuing after 
the breiak up of the continents. 

(4) Centres of origin 

The centre of origin of a group is often regarded as that area having the greatest concentration of 
species. Hale used this method in assessing the centre of origin of the genera Relicina (Hale, 
1975), Pseudoparmelia (Hale, 1976a), Parmelina (Hale, 19766), and Bulbothrix (Hale, 1976c). 

If the numbers of Ramalina taxa occurring in each landmass that once formed Gondwanaland 
are estimated (and these numbers must be only approximate as without recent revisions many 
names could be synonyms or misidentifications) then it is found that 41 Ramahnae occur in 
South America, 42 in Africa, 36 in India, and 30 in Australasia. 

South America and Africa (once West Gondwanaland) appear to have the largest number of 
RamaUnae and could be regarded as the centre of origin of the southern hemisphere taxa. 

The occurrence of high numbers of taxa within the one genus, in a particular region, can, 
however be due to a re-radiation of species after the original dispersal of the taxa. When an 
environment is free of competitive species, newly evolved taxa belonging to the genus will 
become established. This situation has been found in Western Australia amongst the eucalypts, 
where secondary radiation of species has occurred in a region where there is little competition 
from other genera (Pryor, 1981). 

Such a situation may have arisen in Australia with the evolution of the mangrove flora in this 
region (Specht, 19816). Mangroves are the major phorophytes for the present day R. subfrax- 
inea complex. It is proposed that the ancient stock of the R. subfraxinea group could have 
entered Australia via the northern route from East Africa (as this group is regarded as a 
megatherm response group) and become estabUshed along the Australian tropical coastline on 
the newly evolved mangroves. The occurrence of all six chemotypes in the region around 
latitude 18°S indicates an area of speciation from where secondary radiation could have spread 
to the east and west. 

(5) Vicariant taxa 

The Croizat theory of vicariance biogeography (Croizat, 1952; Croizat, Nelson & Rosen, 1974) 
postulates that species which are ecologically similar but occur on different land masses have a 
common ancestry. They are regarded as disjunct populations of once continuously distributed 
taxa which have undergone speciation as physical disruption occurred and Croizat cited many 
Gondwanaland taxa to support his theory. Galloway (1979) employed the same ideas in his 
comments on the distribution of species in the genera Usnea, Pseudocyphellaria, Psoroma, and 
Stereocaulon. 

A number of taxa belonging to the Fistularia group in Australia appear to be part of the 
vicarant taxa of Gondwanaland origin. For example the Australian taxon R. inflata subsp. 
perpusilla has an ecotype which grows on mangroves and closely resembles the East African 
species R. consanguinea which also occurs on mangroves with the main difference in the spore 
size; in some habitats in AustraUa R. inflata subsp. inflata morphology closely resembles some 
forms of the East African taxon R. calcarata, but their chemistry differs in the minor 
constituents. 



RAMALINA IN AUSTRALIA 147 

Chemical difference is the only character which separates the Australian taxon R. filicaulis 
from the American taxon R. anceps. Material similar to the Australian material, and also lacking 
medullary acids, has been cited from South Africa. It is proposed that all three continents 
possess species which could be regarded as vicariant taxa belonging to the R. usnea complex, as 
they are ecologically and morphologically similar. 

On the basis of the evidence discussed above it is postulated that Australia has been populated 
by species of Ramalina since the late Cretaceous. Since west Gondwanaland is accepted as the 
centre of origin for the angiosperms, based on fossil evidence, there is the possibility of these 
acting as phorophytes for ancestral taxa of the present day Ramalina, as that same region is a 
likely centre of origin for Ramalina. The step by step spread of the plant communities would 
have enabled the Ramalinae to spread by one of two migration routes either (1) the southern 
route from South America (after Africa had separated) via the Scotia arc, Antarctica to 
Tasmania and Australia or (2) the northern migration route from East Africa via Madagascar, 
the Mascarenes, Indo-Malaya to northern Australia, by which means the megathermic Ramali- 
nae could have reached tropical Australia. Once established in Australia the megatherm 
Ramalina taxa remained a maritime group as inland habitats would have been inhospitable. 
Secondary radiation from Australia by some of the megathermic taxa appears to have occurred 
with the spread of species to the Pacific and Indian ocean islands. 

The microtherm Ramalinae became established in the mountainous region of southern 
Australia since these habits would have suited their thermal response patterns as Australia 
drifted northwards and the temperatures increased. 

The climatic changes that occurred in the late Miocene Period caused the desiccation of the 
continent. Subsequent speciation of mesotherm Ramalinae could have occurred producing taxa 
which occupied the new and dryer habitats of the south-west and west of Australia. 

In this way the present distribution pattern of the AustraUan Ramalinae reflects their 
palaeobiogeographic and palaeoecological history. 

Taxonomy 

The dehmitation of Ramalina species proved difficult owing to the amount of morphological 
variation shown by each taxon. A large amount of material had to be examined in order to 
ascertain what concepts should be appUed in the delimitation of each taxon. Three taxonomic 
ranks have been used. 

Taxonomic ranks 

(1) Species: The rank of species has been given only to those taxa which can be recognized by 
several distinct morphological traits which collectively produce a particular thallus type (allow- 
ing for an appropriate amount of variation), together with a set of specific anatomical and 
chemical properties, and which show a distinct habitat and distribution range. 

Difficulties arose when two or more chemotypes produced a single morphotype; this problem 
had to be resolved by considering the habitat and distribution range of the taxa, and if the only 
difference found was in the chemistry then these chemical races were treated as conspecific. 

The concept of species pairs (Artenpaare), as proposed by Poelt (1970, 1972) is that species 
which reproduce by vegetative methods (secondary species) have been derived from existing or 
extinct fertile species (primary species), and that primary and secondary species have identical 
(or closely related) chemical components. This view is held by most Uchenologists and species 
pairs have been found in many genera e.g. in the Physcia and allied genera (Moberg, 1977), 
Parmelia sect. Hypotrachyna (Culberson & Hale, 1973), Dirina (Tehler, 1983), and to a lesser 
extent in Alectoria (Brodo & Hawksworth, 1977). 

Only a few records of species pairs have been found in the RamaUnae. Rundel & Bowler 
(1976) discussed R. leptocarpha and R. sublectocarpha as a fertile-sorediate species pair and 
Krog & 0sthagen (1980) have commented that the fertile species R. implectens may be regarded 
as the parent morph of the sorediate species R. farinacea. 



148 G. NELL STEVENS 

None of the Australian sorediate Ramalinae is regarded as derived from any of the fertile taxa 
found in Australia today. 

(ii) Subspecies: The delimitation of a taxon based on well marked discontinuities in characters 
was not always possible. Some Ramalina taxa were found to intergrade into each other, differing 
only subtly morphologically yet extending over large areas. When such complexes were 
examined carefully the most obvious morphological changes were found to coincide with 
different geographical regions. 

The term subspecies has therefore been applied to a taxon which could be shown to occupy a 
particular region within the total distribution range of a large complex (as defined by Hawks- 
worth, 1976). This taxon can be extremely variable within its geographical confines and can 
produce several ecotypes which occupy separate habitats. 

As subspecies intergrade into each other, their delimitation is much more arbitrary than that 
of a species, and depends more on geographical divisions than on chemical or morphological 
differences. Imshaug & Brodo (1966) pointed out that the use of the rank of subspecies 
permitted a classification which is not only practical but one which indicates a possible 
phylogenetic system. The evolutionary relationships that may exist in some of the AustraUan 
taxa have been indicated by the use of the rank of subspecies. 

(iii) Variety: The rank of variety has been used when more or less morphologically identical 
taxa have been found to produce different chemistries which correlate with distinctive distri- 
bution patterns. 

The production of either divaricatic acid or sekikaic acid in populations of the one taxon posed 
a nomenclatural problem. Several Uchenologists have mentioned the same difficulty e.g. in the 
genus Dirinaria (J0rgensen, 1974) and in the genus Ramalina (Imshaug, 1972; Rundel, 1978fl). 
The solution has been to regard both acids as representing chemical races within the one taxon. 

Hybridization 

It is not easy to estabhsh the genetic homogeneity or heterogenity of lichenized fungi. Jahns 
(1974) pointed out that thallus or hyphal fusions do occur in lichens, which could enable a form 
of hybridization to take place. A specimen of R. leiodea recorded by Elix (1982) as being a 
'schizophrenic thallus', had 'left-hand branches containing one acid, the right-hand branches 
another and the central branches containing a mixture of both'. In this case the two acids 
involved were two depsides, not closely related (cryptochlorophaeic acid and boninic acid). 
Another such combination of two acids in one specimen belonging to the R. subfraxinea 
complex has been found during the present study; the specimen was collected at Hinchinbrook I. 
where all acid races of the R. subfraxinea complex are present. The thallus concerned contained 
both boninic acid and salazinic acid, one a depside and the other a depsidone. As these two acids 
are not biosequential metabolites it appears likely that this is a similar form of hybridization to 
that found by Elix (1982), and that both could be regarded as 'chimeras' produced by the fusion 
of hyphae from two different spore types. It is not known what chemistry the thalli produced 
from spores of this plant would have. No other thalli with this combination of acids has been 
found. 

The occurrence of sekikaic acid in the divaricatic acid race ofR. subfraxinea var. subfraxinea is 
regarded as different from the above. Sekikaic acid and divaricatic acid are reasonably closely 
related biosequentially and apparently can interconvert (Elix in Htt.) which would result in 
chemical combinants if the conversion was incomplete. However the hypothesis of interspecific 
hybridization (Brodo, 1978) remains feasible and has to be considered beside the hypothesis that 
incomplete conversion of a precursor acid occurs during the biosynthetic steps from one acid to 
another (Bowler & Rundel, 1978). If hybridization is accepted as the cause of this mixture of 
acids then it would be natural to expect the proportions of each acid to be equal in the thallus. 
However, this was not the case with the taxa examined; in all instances divaricatic acid occurred 
in far larger quantities than did the sekikaic acid. It is therefore suggested that incomplete 
conversion of one acid to another has taken place. 



RAMALINA IN AUSTRALIA 

Table 8 Taxonomically important characters in the Australian Ramatina taxa. 



149 



















major acid constituents 


















orcinol depsides 




(3-orcinol 


























depsidones 






substrate 


thallus 




para- 




meta- 
























































M 










Ramatina species 


O 

8 
8 


o 

8 


3 
1 


73 




% 
(J 

•c 

CO 

> 


o 

'S 
> 


o 

M 
M 

(/3 


u 








'S 


salazinic 
norstictic 




e 

2 





CO 

'S 


R. australiensis 


X 


X 


X 




















X 


R. caespitella 




X 


X 




X 






X 












R. canadensis 


X 




X 




X 


X 
















R. exiguella 


X 




X 




















X 


R. filicaulis 


X 




X 




















X 


R. fimbriata 




X 




X 


X 


X 
















R. glaucescens 


X 




X 






X 




X 












R. litorea 




X 


X 








X 














R. peruviana 


X 




X 




X 






X 












R. reducta 


X 




X 




X 














X 




R. tenella 


X 




X 




X 












X 






R. unilateralis 


X 


X 


X 




X 


X 
















R. whinrayi 




X 


X 






X 




X 












Species complexes 




























R. celastri 




























subsp. celastri 


X 


X 


X 




















X 


subsp. ovalis 


X 




X 




















X 


R. inflata 




























subsp. inflata 


X 






X 




X 
















subsp. perpusilla 


X 






X 




X 




X 












subsp. aus trails 


X 


X 




X 




X 




X 












R. fissa 


X 


X 




X 














X 






R. nervulosa 




























var. nervulosa 


X 




X 




X 


X 
















var. luciae 


X 




X 




X 






X 












var. dumeticola 


X 




X 




X 






X 












R. pacifica 


X 


X 


X 




X 












X 






R. subfraxinea 




























var. subfraxinea 


X 




X 






X 




X 












var. leiodea 


X 




X 














X 








var. confirmata 


X 


X 


X 












X 










var. norstictica 


X 




X 
















X 






R. tropica 


X 


X 


X 
















X 







Description of the genus Ramalina 

Ramalina Ach., Lichenogr. Univ.: 122,598 (1810), nom. cons. 

Thallus fruticose, greenish-grey, pale green or greenish-yellow, caespitose, erect, subpendulous 
to pendulous; branching dichotomous, subdichotomous to irregular or dense intricate; branches 
few to many, twig-like, strap-like to palmate, flat, subterete or terete, solid or inflated; apices 
forked, attenuate, rounded; surface shiny or matt, smooth or rugose, pseudocyphellate; 
holdfast delimited or dispersed; soralia small and punctiform to large and round or elongate; 



150 G. NELL STEVENS 

cortex composed of two layers, the outer cortex hyphae more or less anticlinal or randomly 
oriented, the inner cortex hyphae perichnally arranged; hyphae surrounded by a large amount 
of matrix; medullary hyphae loosely interwoven between the upper and lower cortex or confined 
to the algal zone. 

Apothecia terminal, subterminal, marginal, lateral, laminal; convex, plane or concave, 
sometimes incised; disc pale to flesh coloured; margin concolourous with thallus; asci clavate, 
containing 8 spores; spores hyaline, oblong, ellipsoid or fusiform, straight or curved, 1-septate. 
Pycnidia inconspicuous, pale, usually rare in Austrahan species. 

The algae are Trebouxioid, globose, forming an irregular ring along the edge of the inner 
cortex; when confined to the upper side only the thallus becomes dichroic with a white lower 
surface. 
Type species (conserved): Ramalina fraxinea (L.) Ach. 

Enumeration of the Australian taxa 

The Australian Ramalinae comprise 28 taxa which can be grouped into 17 fertile and 11 
sorediate taxa. Sixteen species have no subspecific taxa recognized, and four species contain a 
total of 12 subspecific taxa. 

Table 8 lists the taxonomically important attributes of each of the 28 taxa; those belonging to 
species complexes are placed after the alphabetically arranged single species, as is also done in 
the description of the taxa. The disposition of the other Ramalina taxa recorded by Wetmore 
(1963) and Weber & Wetmore (1972) is shown in Appendix 1. 

The specimens examined are cited with location, latitude and longitude in degrees, type of 
substrate, date, the collector's name and the herbarium abbreviation (Holmgren, Keuken & 
Schofield, 1981). Collectors initials have been used for those collectors whose specimens are 
cited most frequently in this work. These names have been abbreviated as follows: 



AA 


A. Archer 


ED 


E. Dahl 


GB 


G. C. Bratt 


GK 


G. Kantvilas 


GNS 


G. N. Stevens 


HS 


H. Streimann 


JAE 


J. A. Elix 



JC 


J. Cashin 


JW 


J. S. Whinray 


NCS 


N. C. Stevens 


NS 


N. Sammy 


RF 


R. B. Filson 


RS 


R. D. Seppelt 


RWR 


R. W. Rogers 



Only one specimen is cited from each location for most of the taxa; where the taxon grows 
profusely over a wide area, only specimens from selected sites are fisted: viz, R. celastri subsp. 
celastri, R. celastri subsp. ovalis, R. inflata subsp. inflata, R. inflata subsp. perpusilla, R. inflata 
subsp. australis, and R. fissa. 

Key to Ramalina in Australia 

la. Thallus inflated (partially or totally) 35 

lb. Thallus not inflated 2 

Plant saxicolous 3 

Plant corticolous 12 

Thallus sorediate 4 

Thallus not sorediate 5 

Soredia eruption through eroded lower surface and at apices 6. R. fimbriata (p. 161) 

Soredia contained in well defined soralia, which are marginal and sometimes laminal 

2. R. caespHella (p. 155) 

Thallus made up of a few loose branches 6 

Thallus composed of numerous branches forming a dense cushion 8 

Branches compressed, flat (apothecia marginal) 14. R. celastri subsp. celastri (p. 180) 

Branches subterete to terete 7 

Branching dense, distal, K- (no medullary acids) I. R. australiensis (p. 152) 

Branching sparse, K+ red (salazinic acid) 20. U. tropica (p. 210) 



2a 
2b. 


(lb). 


3a 
3b. 


(2a). 


4a 
4b. 


(3a). 


5a 
5b. 


(3b). 


6a 
6b. 


(5a). 


7a 
7b. 


(6b). 



RAMALINA IN AUSTRALIA 151 

8a (5b). Branches solid 9 

8b. Branches inflated , often split 10 

9a (8a). Apothecia terminal, large 3-10 mm diam 13. R. whinrayi (p. 179) 

9b. Apothecia subterminal (subtending narrow pointed branchlets, apothecia 1-2 mm 

diam.) 8. R.ntorea{p. 169) 

10a (8b). Thallus showing some inflation; splitting and tattering of branches common 11 

10b. Thallus not inflated 12 

11a (10a). Medulla K+ red (salazinic acid) 19. R.fissa{p. 193) 

lib. Medulla K- or K+ pink (divaricatic acid or sekikaic acid) 

15. R. inflata subsp. australis (p. 191) 

12a (2b). Thallus sorediate 13 

12b. Thallus not sorediate 21 

13a (12a). Lower surface of anastomosing strands resting on the medulla 10. /?. reducto (p. 175) 

13b. Lower surface not of anastomosing strands only 14 

14a (13b). Branches few, usually broad at base (soft textured, soralia apical and marginal) 

3. R. canariensis (p. 156) 
14b. Branches numerous, narrow 15 

15a (14b). Branches usually cartilaginous (splitting open at intervals along their length) 

12. R. unilateralis (p. 177) 
15b. Branches not cartilaginous, complete 16 

16a (15b). Medulla K+ red (salazinic acid) 17 

16b. MeduUaK-orK+pink 18 

17a (16a). SoraUa punctiform, mainly apical on branchlets (thallus up to 3 cm high) 

11. U. tenella (p. 176) 
17b. Soralia ellipsoidal or round, marginal on elongate branches (thallus usually 4-8 cm 

long) 18. R.pacifica(p. 201) 

18a (16b). Soralia ellipsoidal or round, marginal on elongate branches 19 

18b. Soralia punctiform, marginal and apical (branching dense, intricate, thallus resilient in 

texture) 9. R. peruviana (p. 170) 

19a (18b). Medulla K+ pink (sekikaic aggregate acids) 20 

19b. Medulla K- (divaricatic acid and stenosporic acid) 17. R. nervulosa var. nervulosa (p. 195) 

20a (19a). Sekikaic aggregate acids ± faint ramalinolic acid 17. R. nervulosa var. dumeticola (p. 197) 
20b. Sekikaic aggregate acids ± faint homosekikaic acid .... 17. R. nervulosa var. luciae (p. 199) 

21a (12b). MeduUaK+red 22 

21b. MeduUaK-orK+pink 23 

22a (21a). Branches flat, sometimes canaliculate, containing norstictic acid (spores 12-16 x 4-5 

^m) 19. R. subfraxinea var. norstictica (p. 208) 

22b. Branches usually subterete, never canaliculate, containing salazinic acid (spores 14-22 

x4-6/>tm) 20. R. tropica (p. 210) 

23a (2 1 b) . Branches subterete to terete , containing no medullary acids 24 

23b. Branches flat, with or without medullary acids 28 

24a (23a). MeduUaK- 25 

24b. MeduUaK+pink 30 

25a (24a). Apothecia lateral and subterminal with branch apex appearing as a long spur (some- 
times black tipped) 4. R. exiguella (p. 158) 

25b. Apothecia never subterminal nor bearing a spur 26 

26a (25b). Distal branchlets occur on otherwise unbranched primary branches 

1. R. australiensis (p. 152) 
26b. Dense distal branchlets not present 27 

27a (26b). Thallus thread-like (with dichotomous branching) 5. R. filicaulis (p. 160) 

27b. Thallus fruticose 28 

28a (27b) . Apothecia marginal on both edges (branches may be canaliculate , no medullary acids) 

14. R. celastri subsp. celastri (p. 180) 



152 G. NELL STEVENS 

28b. Apothecia not marginal on both edges 29 

29a (28b). Apothecia laminal, numerous, and equal in size 14. R. celastri subsp. ovalis (p. 184) 

29b. Apothecia not laminal and numerous and equal in size 30 

30a (29b) . Branches broad with apices forked or lacerated , never simple and attenuate [apothecia 
in axils of branches (up to 5 mm diam.) or terminal on the broad margins of the 
apices, or small and sparsely spread laminally] 7. R. glaucescens (p. 163) 

30b. Branches usually narrow with apices simple and attenuate or bent to form a spur below 

apothecium 31 

31a (30b). Branches narrow with dichotomous branching (apothecia marginal or subterminal or 

intheaxilsofbranches,small0-5-l-5mmdiam.) 7. R. glaucescens (p. 163) 

31b. Branches narrow to broad, branching usually from the base only 32 

32a (31b). MedullaK+pink 33 

32b. MedullaK- 34 

33a (32b). Branches elongate, narrow, often tuberculate, containing sekikaic acid 

19. R. subfraxinea var. subfraxinea (p. 203) 

33b. Branches not elongate, rarely tuberculate, containing cryptochlorophaeic acid 

19. R. subfraxinea var. confirmata (p. 205) 

34a (32b) . Thallus containing boninic acid 19. R. subfraxinea var. leiodea (p. 207) 

34b. Thallus containing divaricatic acid 19. R. subfraxinea var. subfraxinea (p. 203) 

35a (la). Thallus sorediate 6. R. Bmbriata (p. 161) 

35b. Thallus not sorediate 36 

36a (35b). Thallus button-like, bearing apothecia (3 mm diam.) between sterile branches or 
totally sterile with eroded apices, medullary reaction K-, saxicolous 

15. R. inttata subsp. australis (p. 191) 
36b . Thallus not button-like nor bearing apothecia between sterile branches 37 

37a (36b). Branches split open and tattered 38 

37b. Branches perforate but complete 39 

38a (37a). Thallus grossly inflated, few or no perforations, splitting open of branches, medullary 

acids divaricatic or sekikaic 15. I?, inffata subsp. australis (p. 191) 

38b. Thallus grossly inflated or split apart, no perforations, medullary acid salazinic 

19. R. fissa (p. 193) 
39a (37b). Thallus semi-inflated to almost compressed, apothecia never concave 

15. R. inttata subsp. perpusilla (p. 188) 
39b. Thallus totally inflated, apothecia concave and innate 15. R. inflata subsp. inffata(p. 185) 

1. Ramalina australiensis Ny\. Plate 7, fig. 1; Plate 13, fig. 1. 

in Bull. Soc. Linn. Normandie II, 4: 120 (1870). Type: Australia, West Australia, Swan River, 1846, 

Verreaux s.n. (PC! - holotype; H-NYL 37491! - isotype; usnic acid). 
Ramalina myrioclada Miill. Arg. in Flora, Jena 66: 20 (1883). Type: Australia, New South Wales, Twofold 

Bay, 1882, White s.n. (G! - holotype; usnic acid). Plate 13, fig. 2. 

Thallus corticolous, rarely saxicolous, pale green to grey-green, erect to pendulous, 3-10 cm 
long, exceptionally to 20 cm; branching sparse in the basal region of the primary branches, 
becoming dense and irregular towards the apices where small branchlets form tufts; branch 
width 0-2-0-5(-l -0) mm, primary branches usually subterete and fine, or thick, compressed and 
coarse, branches never tortulose, often subsidiary short branchlets occur at right angles to the 
primary branch, apices of branchlets often hooked; surface matt, smooth or furrowed, longi- 
tudinal splitting occurs on the surface in coarser forms; pseudocyphellae often present; holdfast 
delimited although several branches arise from it, attachment of branches to the substrate at 
intervals along their length may occur; soralia absent. 

Apothecia uncommon, lateral on the primary branches, usually towards the base; disc 

Plate 7 Thallus growth forms in the Australian Ramalinae. Scale in mm. Fig. 1 R. australiensis. Fig. 2 R. 
caespitella. Fig. 3 R. canariensis. Fig. 4 R. canariensis (large form). Fig. 5 R. exiguella. Fig. 6 R. filicaulis. 
Fig. 7 R. fimbriata. Fig. 8 R. litorea. ► 



RAMALINA IN AUSTRALIA 



153 




lllllillllllllllllllllllllliiliillllilllllllllllllllllllllillllllililililiililllliill til 



iiiiiii 




iifliii tilllllllllllllllllllllllill 



154 



G. NELL STEVENS 



0-5-1 •0(-l -2) mm diam., plane to convex, margin entire; spores ellipsoid to fusiform, straight, 
12-16(-20) X 4-6 fxm (Plate 4, fig. 2). 

Chemistry. Usnic acid only. 

Remarks. The name R. australiensis as the oldest name has been upheld for this taxon, 
overriding the more commonly used name R. myrioclada. However the location given for the 
type material - Swan River, Western Australia - is dubious. All recent collections have been 
made in eastern Australia with no other material being found in the west. Whether the original 
material from Western Australia was wrongly recorded was not able to be ascertained. 

The type material of R. australiensis has branchlets with black tips but this feature (as with R. 
exiguella) is not present in most specimens. The type is a coarse thallus form of this taxon, rather 
resembling that of the subtropical Queensland collection. There is not sufficient type material to 
obtain an overall picture of the total thallus. 

The degree of secondary branching varies from specimen to specimen, those with dense 
branching resemble R. peruviana whereas others have sparse branching producing an open 
thallus form resembling the morphology of R. filicaulis N. Stevens. 

The names R. usneoides and R. usnea have been applied to some specimens oiR. australiensis 
collected in New South Wales but the Australian taxon can be distinguished by its branching 
pattern and subterete branches instead of the characteristic flattened branches of R. usnea 
sens. str. 

Distribution and habitat. This subtropical/warm temperate maritime species does not have a 




Fig. 8 Distribution of R. australiensis • and R. filicaulis * . 



RAMALINA IN AUSTRALIA 155 

wide distribution (Fig. 8). It occurs along the coast from Yamba 29°S to Twofold Bay 37°S and 
has a disjunct occurrence on the Queensland coast at Toogoom (25°S). This Queensland 
occurrence cannot be adequately explained, although the rainfall in this area is similar to that 
which occurs along the New South Wales coastline. The Queensland material is extremely 
coarse, it was collected from Callitris columellaris which was growing on exposed sand dunes. 
The morphology of several Ramalina species collected from this area was also extremely coarse, 
indicating environmental adaptation. 

The major phorophyte of R. australiemi is Avicennia marina, which grows in sheltered 
positions along the bays and rivers of the New South Wales coastline. However this Ramalina 
has also been collected from rocks at Port Hacking and Port Macquarie, which indicates it is not 
substrate specific. 

This taxon occurs on the offshore islands of the North Island of New Zealand which lie at 
equivalent latitudes to those where R. australiensis occurs in Australia. The species has not been 
recorded anywhere else. 

World distribution: Australia and New Zealand. 

Queensland: Toogoom, 25°15'S, 152°40'E, on Callitris columellaris, 1983, GNS (BRIU4217NS). New 
South Wales: Yamba, 29°27'S, 153°20'E, on Avicennia marina, 1977, GNS (BRIU 1978NS); Nambucca 
Hds. 30°41'S, 153°00'E, on Avicennia marina, 1977, GNS (BRIU1998NS); Port Macquarie, 31°27'S, 
152°55'E, on metamorphic rock, 1975, J AE (JAE 1081); Patonga 33°30'S, 151°20'E, on Avicennia marina, 
1978, GNS (BRIU2314NS); Gosford, 33°30'S, 151°20'E, 1978, JAE (JAE 4715); Newport, 33°40'S, 
151°20'E, on mangroves, 1888, F. R. M. Wilson (MEL 9409); Lilli Pilli Beach, 34°04'S, ISriO'E, on 
sandstone, 1975, JAE (JAE 1181); Sussex Inlet, 35°10'S, 150°35'E, on Avicenna marina, 1977, RWR 
(BRIU1902NS); Buckenbowra R. 35°42'S, 150°06'E, on Avicennia marina 1983, HS (CBG8306040); 
Shoalhaven R. 34°52'S, 150°42'E, on Aegiceras corniculatum , 1975, P. 5ae«ger (BRIU2137NS). 

2. Ramalina caespitella N. Stevens Plate 7, fig. 2. 

in Lichenologist 18: 183 (1986). Type: Australia, Long Island (Bass Strait), 1969, Whinray s.n. (MEL! - 
holotype; sekikaic acid, 4'-0-demethylsekikaic acid, ± several terpenes, and usnic acid). 

Thallus saxicolous, pale greenish-grey to stramineous, minute, caespitose, rigid, up to 1-0 cm 
high; branching subdichotomous, irregular, narrow thalli densely branched, broad thalU 
sparsely branched; branch width 0- 1-2-0 mm, exceptions to 6 mm, branches solid, flat to 
subterete, narrow branches nodular, apices broad and blunt; surface shiny; holdfast delimited or 
diffuse, soraUa marginal and laminal, round to ellipsoidal, becoming fissural on the upper 
surface occurring on eroded areas of the lower surface and at the apices. Apothecia rare, laminal 
near the branch apices, rarely terminal, disc 1-0 mm diam, concave, margin thick, entire; spores 
not produced. 

Chemistry: sekikaic acid (M), 4'-0-demethylsekikaic acid (t), ± several terpenes and usnic acid. 

Remarks. Ramalina caespitella produces two morphotypes which intergrade into one another. 
One morph has narrow subterete to terete nodular branches with apical and marginal soralia, 
whilst the other morph produces very thick, flat to subterete branches bearing numerous soralia 
on both upper and lower surfaces. The extreme button-like morphology of both forms is 
indicative of their exposed, saxicolous habitat. No other taxon has been found to have a close 
affinity with R. caespitella. 

Distribution and habitat. This temperate maritime taxon is found on several of the islands in 
Bass Strait and along parts of the Tasmanian coastline (Fig. 9). It occurs on siliceous rocks (e.g. 
granite on the islands and on quartzite or dolerite at Tasmanian locations) and often occupies 
cliff -face habitats. 

World distribution. Australia. 

Bass Strait: Hogan Group: Hogans I, 39°13'S, 146°59'E, on granite, 1973, JW (MEL1013070); Kents 
Group, Dover I, 39°29'S, 147°17'E, on granite, 1970, yW(MEL1012702); Deal I, 39°30'S, 147°22'E, on 
granite, 1972,/W(MEL1012940); Furneaux Group: West Sister 1, 39°45'S, 147°56'E, on granite, 1966, JW 



156 



G. NELL STEVENS 




Fig. 9 Distribution of R. caespitella. 

(MEL1026132); Cape Barren I, 40°25'S, 148°10'E, on granite, 1969, JW (MEL1018083); Big Chalky I, 
40°06'S, 147°54'E, on granite, 1972,/W(MEL1018053); Isabella Reef, 40°25'S, 148°30'E, on granite 1973, 
JW (MEL522220); Flinders I, 39°45'S, 147°56'E, on rock, 1965, RF (MEL1026188); Swan I, 40°41'S, 
148°05'E, on rock, 1974,/W(MEL1019119); Tasmania: West Point, 40°55'S,145°15'E, on quartzite, 1981, 
GK (362/81); Rocky Cape, 40°51'S, 145°31'E, on quartzite, 1970, GB (H032637); Eddystone Pt., 41°00'S, 
148°23'E, on granite, 1973, GB (H032640); Arthurs River, 41°07'S, 144°42'E, on quartzite, 1970, GB 
(H032639); Bruny I., 43°29'S, 147°09'E, on granite, 1973, GB (H032636). 



3. Ramalina canariensis Steiner 



Plate 7, figs 3-4. 
isotype; 



in Ost. bot. Z. 9: 8 (1904). Type: Gran Canaria, Tafira, Bornmtiller, PI. Canar. 3500 (W! 
divaricatic acid and usnic acid). 

Thallus Gorticolous, pale green to grey green, erect to subpendulous, l-3(-5) cm long; 
branching palmate or irregular; branch width l-3(-10) mm, apices blunt, usually split with 
margins separated; surface matt, smooth to rugose, sometimes coarsely reticulately ridged by 
chondroid strands, pitted distally; holdfast delimited; soralia marginal and apical occurring 
through the separation of the upper and lower surfaces. 

Apothecia not seen. Reported as rare. 

Chemistry. Divaricatic acid, usnic acid. 

Remarks. Fine thalli of this taxon come close to resembling immature thalli of R. unilateralis 



RAMALINA IN AUSTRALIA 



157 



and both species contain divaricatic acid, but the cartilaginous surface and the finely lacerated 
apices of R. unilateralis enable the two to be separated. 

Distribution and habitat. The distribution pattern of R. canadensis in Australia is mainly 
maritime in areas experiencing a Mediterranean type of climate (Fig. 10). The only inland 
collections are two South Australian locations (Springton and Kuitpo Forest) at elevations of 
300 m where frequent mists occur. 

Krog & 0sthagen (1980) mentioned that in the Canary Islands (28°N) this species 'seems to 
prefer well lit sites with a certain influence of mist . . . between 400 and 1000 m altitude' . In some 
of the drier Australian coastal sites morning fogs are prevalent. 

Along the southern coastline of AustraUa this species grows in well-lit situations protected 
from winds, usually on twigs of Leucopogon parviflora, Casuarina, and Melaleuca spp. It has 
also been found on mangroves {Avicennia marina) along the eastern coastline south from 
latitude 35°S. Ramalina canariensis occurs in homogeneous colonies or immixed with R. fissa. 

Overseas specimens were examined from South Africa (Cape Province, latitude 33°S at 800 m 
elevation) and Chile (Valparaiso, latitude 33°S at sea level, on dune shrubs). Both of these 
regions have climates similar to southern Australia and other Mediterranean regions. Ramalina 
canariensis was previously thought to grow only in the northern hemisphere in the Mediterra- 
nean region and on the Canary Islands. 

World distribution. Mediterranean region, Canary Islands, South Africa, Australia, and South 
America. 





120 




130 


iio 






iio 









800km 


isa'^ 


f& 


1 






10> 






/" 


V 


C 


^ 








-- 


V 


20^ 












"' \~~~~ 


"^^ 


^ /* 


Z^"^*" 


1 







J 


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3(K 
40> 



Fig. 10 Distribution of /?. canariensis. 



158 G. NELL STEVENS 

New South Wales: Sussex Inlet, 35°10'S, 150°35'E, on Avicennia marina, 1977, RWR (BRIU1903NS); 
Buckenbowra R. 35°42'S, 150°06'E, on Avicennia marina, 1983, HS (CBG8306039); Batemans Bay, 
35°45'S, \5(fWB on Avicennia marina, 1978, GN5(BRIU2171NS);Batehaven Beach, 35°45'S, 150°13'E, 
onCa5uarmflsp.,1975,y>l£(JAE1061);BurrenwarraPt.,35°49'S, 150°14'E,onCfl5Mflnflsp.,1975,ME 
(JAE 1246). Victoria: Cape Conran, 37°48'S, 148°44'E, on Leucopogon sp., 1983, GNS (BRIU3701NS); 
Westernport Bay, 38°13'S, 145°18'E, on Avicennia marina, 1978, GNS (BRIU2336NS); Gunnamatta 
Beach, 38°21'S, 144°45'E, on Monotoca elliptica, 1983, GNS (BRIU3692NS); Warrnambool, 38°23'S, 
142°29'E, 1886, F. R. M. Wilson (NSW.L4311); Portland, 38°20'S, 141°36'E, 1896, F. R. M. Wilson (MEL 
9419); Portsea, 38°25'S, 144°42'E, on tree on sea cliff, 1964, /. Williams (MEL1026126); Somers, 38°24'S, 
145°10'E, 1964, Marie (MEL1026173). South Australia: Kuitpo forest, 35°07'S, 139°16'E, on Pinus sp., 
1968, RWR (BRIU1426NS); Kangaroo I. Lockwood Corner, 35°39'S, 137°38'E, 1967, G. Jackson 
(S. A. 97647604); Mt Lofty Ra., Carey Gully, 34°45', 139°00'E, on Exocarpus cupressiformis, 1976, JAE 
(JAE 2853); Springton, 34°45'S, 139°05'E, on Callitris sp., 1976. JAE (JAE 2807); Eyre Peninsula: 
Memory Cove, 34°40'S, 135°50'E,onMe/fl/eMCflsp., 1970,i?F(MEL1018638);Elliston,32°59'S, 134°30'E, 
1967, N. Donner (SA97648163). Western Australia: Gairdner River, 34°14'S, 119°17'E, on Hakea sp., 
1980, D. Richardson (WA000943); Recherche Arch. Long I, 34°03'S, 121°57'E, on Melaleuca sp., 1950, 
JW (MEL10020). Bass Strait: Furneaux Group: Flinders I, 40°00'S, 148°00'E, on shrub, 1977, JW 
(MEL10225619); East Sister I, on dune shrub, 1972,yW(MEL1025619); West Sister 1, 39°42'S, 147°56'E, 
on Pomaderris apetala, 1969, JW (MEL521577); Hogan Group: Long I, 39°12'S, 147°01'E, on Banksia 
integrifolia, 1973, JW (MEL1012966). 

4. Ramalinaexiguella Stirton Plate?, fig. 5; Plate 13, fig. 3. 

in Trans. Proc. R. Soc. Vict. 17: 68 (1881). Type: Australia, Queensland, Brisbane, Bailey 91 (BRI! - 

holotype; BM - isotype; usnic acid). 
Ramalina pertenuis Magnusson in Ark. Bot. 32A (2): 7 (1945). Type: Oahu, Keeawa awa Valley, 1922, 

Skottsberg s.n. (GB! - part of holotype; usnic acid). 

Thallus corticolous, grey-green, caespitose, erect, rigid, up to 3 cm high; branching mainly 
from the base, squarrose branchlets sometimes present on main branches; branch width 0-3-1-0 
mm, branches subterete, narrow, apices attenuate, sometimes black tipped; cortex shiny, 
longitudinally grooved giving a string-like appearance; pseudocyphellae linear; holdfast delimi- 
ted; soralia absent. 

Apothecia few to many, usually marginal along the branches and/or subterminal causing the 
branch apex to become geniculate with a long, attenuate spur; disc. 0-2-2-0 mm diam. , concave, 
plane to convex; margin entire, thick on immature apothecia; spores ovoid to ellipsoid or 
gibbous and slightly curved, (12-) 14-16 x (5-)6-8 ixm (Plate 5, fig. 1). 

Chemistry. Usnic acid only. 

Remarks. The affinities of this taxon lie with R. gracilis (Pers.) Nyl. ; Nylander (1870) recorded 
material from Brazil, Madagascar, and Australia as belonging to R. gracilis. The type of R. 
gracilis was not available from either L or PC, so any relationship between these two taxa was 
not able to be checked. Some material of R. gracilis held at BM is morphologically similar to R. 
exiguella but contains salazinic acid. Krog & Swinscow (1976) reported that R. attenuata (Pers.) 
Tuck, from Domingo, resembled R. exiguella but contained psoromic acid. Landron (1972) 
stated that R. gracilis contained psoromic acid whereas R. attenuata contained salazinic acid and 
that the latter species is synonymous with R. rigida Ach. Howe (1913-14) placed R. rigida and 
R. gracilis as synonyms of/?, attenuata. 

It could be that this group of taxa is a complex with several chemical races, but until the type 
material of all of these species can be located and examined, the relationship between these taxa 
cannot be resolved. Stirton stated in his type description that R. exiguella had affinities with R. 
melanothrix Laurer, but the latter species is quite distinct and the type material does not 
resemble Australian material oiR. exiguella. 

Distribution and habitat. The distribution of this coastal species is continuous along the eastern 
shoreline of Australia from latitudes 23°S-35°S, with additional occurrences at 18°S and 2rS. Its 
occurrence on the coastal hills is rare, one tropical collection was made from Abbey Peak, 650 m 



RAMALINA IN AUSTRALIA 



159 



elevation at latitude 14°S and one subtropical collection from Mt Mothar 320 m elevation at 
latitude 26°S (Fig. 11). 

Ramalina exiguella grows on several species of mangrove (viz. Rhizophora stylosa, Ceriops 
tagal, Avicennia marina, and Aegiceras corniculatum) and on other coastal trees {Casuarina 
equisitifolia and Callitris columellaris) which grow close to the water. It also occupies very 
exposed situations in the dune communities facing the ocean, where it is often the only lichen 
present. 

This taxon shows a marked difference in size of thallus in different habitats. The thallus may 
be minute (1-5 cm high) and narrow (as the name implies) or extremely coarse and ropy and up 
to 3 cm high. South of 28°S the thalli tend to produce lateral branchlets along the branches. 

In sheltered habitats, R. exiguella grows in association with R. inflata subsp. perpusilla, R. 
pacifica, R. subfraxinea var. leiodea, and R. subfraxinea var. confirmata. 

World distribution. East Africa, Australia, and New Zealand. 

Queensland: Abbey Pk 14°18'S, 144°30'E, on shrubs, 1983, C. McCracken (BRIU4232NS); South Mission 
Beach, 17°55'S, 146°05'E, on Ficus sp., 1983, GNS (BRIU4159NS); Eimeo, 2r06'S, 149°10'E, on Cerios 
tagal, 1979, GNS (BRIU1860NS); Yeppoon, 23°08'S, 150°45'E, on Casuarina sp., 1980, RWR 
(BRIU2221RR); Keppel Sands, 23°21'S, 150°47'E, on Rhizophora stylosa, 1975, RWR (BRIU890RR); 
Rhodds Peninsula, 24°02'S, 151°40'E, on Rhizophora stylosa, 1975, GNS (BRIU1277NS); Turkey, 
24°06'S, 151°37'E, on Ceriops tagal, 1975, GNS (BRIU1156NS); Round Hill Head, 24°10'S, 151°54'E, on 
Aegiceras corniculatum, 1975, RWR (BRIU1935RR); Burnett Heads, 24°45'S, 152°25'E, on Casuarina, 
1976, GNS (BRIU4203NS); Eraser I, Wathumba Ck, 24°50'S, 153°13'E, on Ceriops tagal, 1975, GNS 




Fig. 11 Distribution of/?, exiguella. 



160 G. NELL STEVENS 

(BRIU642NS); Urangan, 25°19'S, 152°55'E, on Rhizophora stylosa, 1975, GNS (BRIU631NS); Boreen 
Point, 26°18'S, 152°58'E, on Callitris columellaris , 1982, GNS (BRIU4181NS); Cooran, 26°17'S, 
152°50'E, on Araucaria cunninghamii, 1985, NCS (BRIU4335NS); Noosa River mouth, 26°24'S, 
153°04'E, on Casuarina sp., 1980, GNS (BRIU3184NS); Maroochydore, Maroochy R, 26°42'S, 153°05'E, 
on Avicennia marina, 1976, GNS (BRIU1313NS); Currimundi Lake, 26°45'S, 153°08'E, on Casuarina sp., 
1974, RWR (BRIU3078RR); Bribie I, 27°02'S, 153°10'E, on Avicennia marina, 1983, GNS 
(BRIU3711NS); Moreton I, Koorigal, 27°20'S, 153°25'E, on Callitris columellaris, 1980, RWR 
(BRIU4179NS); Stradbroke I, Myora Springs, 27°28'S, 153°24'E, on Avicennia marina 1977, GNS 
(BRIU1585NS); Thorneside, Tingalpa Ck 27°28'S, 153°11'E, on Rhizophora stylosa, 1977, GNS 
(BRIU1742NS); Wellington Point, 27°28'S, 153°15'E, on Avicennia marina, 1975, RWR (BRIU5648RR); 
Ormiston, 27°30'S, 153°16'E, on Avicennia marina, 1975, RWR (BRIU5459RR); Coochie Mudlo I, 
27°35'S, 153°20'E, on Callitris columellaris, 1975, GNS (BRIU4579RR); Point Talburpin, 27°35'S, 
153°17'E, on Rhizophora stylosa, 1982, GNS (BRIU4189NS); Little Rocky Point, 27°38'S, 153°18'E, on 
Ceriops tagal, 1977, GNS (BRIU1437NS). New South Wales: Hastings Point, 28°26'S, 153°32'E, on 
Avicennia marina, 1977, GNS (BRIU1929NS); Brunswick Heads, 28°33'S, 153°33'E, on Callitris columel- 
laris, 1982, GNS (BRIU3316NS); Byron Bay, 28°40'S, 153°36'E, on Cupaniopsis sp., 1979, RWR 
(BRIU3179NS); Ballina, 28°54'S, 153°33'E, on Avicennia marina, 1977, GNS (BRIU1961NS); Yamba, 
29°27'S, 153°20'E, on Avicennia marina, 1977, GNS (BRIU2125NS); Woolgoolga, 30°06'S, 153°12'E, on 
dune shrubs, 1982, GNS (BRIU4178NS); Sawtell, Boambee Ck, 30°22'S, 153°06'E, on Avicennia marina, 
1977, GNS (BRIU2034NS); Macksville, 30°41'S, 153°00'E, on Avicennia marina, 1982, GNS 
(BRIU4177NS); Macleay River, 30°54'S, 153°03'E, on Casuarina sp., 1982, GNS (BRIU4175NS); Port 
Macquarie, 31°27'S, 152°54'E, on shrub, 1975, JAE (JAE 1260); Port Macquarie, 31°27'S, 152°54'E, on 
Avicennia marina, 1977, GNS (BRIU1983NS); Laurieton, 31°20'S, 152°55'E, on Avicennia marina, 1977, 
GNS (BRIU4339NS); Taree, Saltwater, 31°54'S, 152°34'E, on tree, 1977, JAE (JAE 4014); Booti Booti, 
32°15'S, 152°31'E, on Casuarina sp., 1982, GNS (BRRJ4332NS); Patonga, 33°30'S, 151°20'E, on 
Avicennia marina, 1978, GNS (BRIU2318NS); Towra Point, 34°03'S, 151°10'E, on Avicennia marina, 
1977, C. Scarlett (BRIU641CS); Batehaven, 35°45'S, 150°10'E, on dead Casuarina sp., 1974, JAE (JAE 
417). 

5. Ramalina fiUcaulisN. Stevens, sp. nov. Plate 7, fig. 6. 

Thallus corticolus, virido-griseus, pendulus vel decumbens usque ad 20 cm longus, ramificatione pro parte 
majore dichotoma; rami lati ad 0-2-0 -5 mm, tenues, filicaules, subterete vel teretes, apice attenuati et 
filamentosi, saepe uncinatus; pagina laevigata, fere striata; soralia nulla. Apothecia rara, lateralia, disco 
piano vel convexo, ad 0-5-1 -0 mm in diametro; sporae ellipsoideae vel fusiformes, rectae vel raro 
curvatae, 12-16(-20) x 4-6 /u.m. Acidum usnicum tantum continens. 

Holotype: Australia, Queensland, Lamington Plateau, 9 km N of O'Reillys in rain-forest on Araucaria 
cunninghamii, alt. 920 m. 27 November 1983, G. N. Stevens 3730 (MEL 1048085!). 

Thallus corticolous, green-grey, pendulous or decumbent, up to 20 cm long; branching 
predominantly dichotomous, curving at the axils; branches width b-2-0-5 mm, narrow, thread- 
like, subterete to terete, tapering, apices attenuate and filamentous, often hooked; surface 
smooth, with linear pseudocyphellae; holdfast small, deUmited but branches often attached to 
the substrate along their length; soralia absent. 

Apothecia uncommon, lateral sessile, disc 0-5-1 -0 mm diam. , plane to convex; margin entire; 
spores ellipsoid to fusiform, straight or rarely curved, 12-16(-20) x 4-6 (xm. 

Chemistry. Usnic acid only. ' 

Remarks. Ramalina filicaulis may prove to be a member of the R. usnea complex because, apart 
from a chemical difference, fine specimens of R. filicaulis (no medullary acids) come close to the 
morphology of the type material of R. anceps (norstictic, ± salazinic acids). Landron (1972) 
recorded R. anceps as a species in the Caribbean Islands (10°N-2rN) that occurs in lower 
montane rain-forest, most abundant on trees along the edge of the forest and in areas of high 
rainfall. 

Rundel (1978a) recorded that the R. usnea race with no medullary acids occurs most 
abundantly in eastern South America (23°S-28°S), a distribution pattern closely resembUng the 
Australian taxon, but the flattened branches of R. usnea do not occur in R. filicaulis. There are 
morphological similarities between the Canary Island species R. chondrina and the Australian 



RAMALINA IN AUSTRALIA 161 

taxon; both are subtropical species which occur in the mountains. Krog & 0sthagen (1980) 
recorded R. chondrina as sparse and scattered on trees in laurel forests between 550 and 900 m 
elevation at 27°30'N to 28°30'N. However, the anatomy of these two species differs; R. 
chondrina has an even cylinder of supportive tissue, whereas R. filicaulis has an uneven, ribbed 
cylinder of supportive tissue. 

Distribution and habitat. This species is rare in occurrence, being known from only a few 
locations (Fig. 8). It occurs profusely at Jimna State Forest (26°40'S) and at Lamington Plateau 
(28°15'S) at elevations between 600-900 m. Two early collections may have come from lower 
elevations in the Rockhampton area (23°S), but little information was available on the 
herbarium specimens. At both Jimna and Lamington this taxon grows almost exclusively on the 
trunks and branches of Araucaria cunninghamii (hoop pine) in the rain-forest. The only other 
phorophyte is Citriobatis pauciflora, a shrub which is a remnant of the rain-forest. 

The climatic data recorded for the occurrence of the hoop pine is also relevant for R. filicaulis. 
Webb & Tracey (1967) record that regions supporting Araucaria cunninghamii have a mean 
annual temperature in the subtropics between 12°-23°C, with occasional frosts; the annual 
average rainfall varies from 800-1800 mm with fogs and dew augmenting precipitation in upland 
areas. Hoop pine occurs as part of the climax forest only on soils of relatively low fertility (e.g. 
Jimna, on metamorphics); on soils of high fertility it does not occur in the climax forest except as 
rare veteran trees (e.g. Lamington Plateau, on basalt). 

Because of the rarity of this Ramalina it is difficult to assess what environmental factors are 
limiting its occurrence elsewhere. 

World distribution. Australia. 

Queensland: Jimna, 26°40'S, 152°28'E, on Araucaria cunninghamii, 1980, RWR (BRIU2202RR); Jimna, 
Marumba View, 26°40'S, 152°28'E, on Araucaria cunninghamii, 1980, GNS (BRIU3349NS); Lamington 
Plateau, O'Reillys, 28°15'S, 153°08'E, windfall, 1970, ED (O); O'Reillys, 28°15'S, 153°08'E, on Citrio- 
batus pauciflora, 1984, P. Merottsy (BRIU4344NS). 

6. Ramalina fimbriata Krog & Swinscow Plate 7, fig. 7. 

in Norw. J. Bot. 21: 117 (1974). Type: Kenya, Central Province, Nyeri District, Krog & Swinscow 
2K36/137 (O! - holotype; BM! - isotype; divaricatic acid and usnic acid). 

Thallus saxicolous, pale green to stramineous, small, caespitose, usually very compact and 
button-Uke, often minute; up to 1-0 cm high; branching dense and intricate, distally producing 
branchlets; branch width 0-3-1 -0 mm, branches flat to subterete, variably inflated, some 
branches with slit-like perforations; surface shiny to matt, smooth; holdfast diffuse; soralia form 
from disintegration of the lower cortex occurring laminally and at the apices (Plate 1, fig. 5). 

Apothecia uncommon, but numerous on some thalU, lateral or terminal, disc 0-5-3-0 mm 
diam., concave, plane or convex; margin entire or incised at maturity; spores ellipsoid, straight 
or rarely curved, 8-13 x 4-5 /am. 

Chemistry. Divaricatic acid (often in trace amounts). 

Remarks. Reduced forms of R. unilateralis come close to the morphology of R. fimbriata, but 
the former species is corticolous in Australia and shows no inflation of the branches. The 
northern hemisphere species R. pollinaria, when growing on rock, produces thalli closely 
resembling R. fimbriata, but the former produces evernic acid. It is suggested that convergence 
of morphology due to similarity of habitats is the reason for the resemblance. 

Distribution and habitat. Ramalina fimbriata has a distribution range from latitude 35°S-43°S 
over most of the Victorian highlands extending into eastern Tasmania (Fig. 12). In AustraUa this 
species is saxicolous only, although in East Africa it is both saxicolous and corticolous (Krog & 
Swinscow, 1974). 

Its occurrence on several types of rock (viz. granite, sandstone, conglomerate, trachyte, and 
dolerite) shows it is not substrate specific. In some localities R. fimbriata occupies sheltered 
overhangs and ledges, habitats which would have little moisture available for the thalli, apart 



162 



G. NELL STEVENS 




Fig. 12 Distribution of R. fimbriata. 



from vapour or water droplets from the overhang. Yet large monotypic populations are found in 
such habitats. Fertile specimens were found at Mt Arapiles under a rock overhang. The 
production of apothecia in this sorediate species is rare and the ascospores were produced only 
sparingly, so that mature spores were hard to find. 

Other habitats where this species occurs are exposed rock surfaces where the small thalli are 
further reduced in size, indicating the harsh environmental conditions prevailing (e.g. cold 
winter temperatures with frosts and/or snow and hot summer temperatures). Krog & Swinscow 
(1974) record R. fimbriata growing on rock in the alpine zone of East Africa at altitudes of 
3200-4100 m. 

The compact button-like thalli commonly collected in Australia are smaller than the cortico- 
lous Kenyan material recorded by Krog & Swinscow (1974), but in all other characteristics they 
are identical. 

World distribution. Africa, Australia, and New Zealand. 

New South Wales: Budawang Ra. 35°22'S, 150°03'E, on'conglomerate, 1981, W. A. Weber (COLO 616); 
Australian Capital Territory: Tharwa, 11 km SW of, 35°31'S, 149°04'E, on granite, 1979, M£ (JAE 6195); 
Booroomba Rocks, 35°32'S, 149°00'E, on granite, 1979, HS (CBG7910804); Mt Clear, 35°43'S, 149°07'E, 
on granite, 1980, JAE (JAE 9013). Victoria: Mt Arapiles, 36°45'S, 141°50'E, on sandstone, 1981, RWR 
(BRIU2420RR); 1969, A. C. Beauglehole (MEL1013299); Melville Caves Park, 36°39'S, 143°42'E, on 
rock, 1964, RF (MEL1516515); Bogong High Plains, 36°44'S, 147°18'E, on rock, 1954, C. Skewes 
(MEL1025567); Mt Ararat, 37°20'S, 142°52'E, c. 1875, D. Sullivan (MEL 9471); Hanging Rock, 37°23'S, 



RAMALINA IN AUSTRALIA 163 

144°37'E, on trachyte, 1976, GNS (BRIU3127NS); Yea, 37°13'S, 145°26'E, on granite, 1964, RF 
(MEL1026128); You Yangs, 38°00'S, 144°29'E, on granite, 1966, GB (H032638). Tasmania: Avoca, 
41°45'S, 147°42'E, on granite, 1972, GB (H032552); Freycinet National Park, Mt Amos, 42°13'S, 
148°18'E on granite, 1979, JAE (JAE 5522); Woods Quoin, 42°16'S, 147°05'E, on dolerite, 1972, GB & 
JC (H032336); Lake Tooms Rd, 42°20'S, 147°28'E, on rock, 1974, GB & JC (H032551). 



7. Ramalina glaucescens Krempelh. Plate 8, figs 1-12; Plate 13, fig. 4. 

in Verh. zool.-bot. Ges. Wien 30: 333 (1880). Type: Australia, Victoria, Mount Ararat, 1875, Sullivan s.n. 

(M! - holotype; sekikaic acid and usnic acid). 
Ramalina leiodea var.fastigiatula Miill. Arg. in Flora, Jena 66: 21 (1883). Type: Australia, Victoria, The 

Grampians, Sullivan 14 (G! - holotype; sekikaic acid (trace) and usnic acid). Plate 13, fig. 6. 
Ramalina lacerata Miill. Arg. in Flora, Jena 66: 20 (1883). Type: Australia, Western Australia, Eucla, 

Oliver s.n. (G! - holotype; M!, MEL!, UPS! - isotypes; no medullary acids usnic acid only). Plate 13, 

fig. 5. 
Ramalina brevis F. Wilson in Victorian Nat. 6: 69 (1889). Type: Australia, Victoria, Warrnambool, 1887, 

F. R. M. Wilson s.n. (NSW! - holotype (L4091); no medullary acids, usnic acid only). Plate 13, fig. 8. 
Ramalina calicaris var. australica Rasanen in Suomal. eldin-ja kasvit. Seur. van. Tiedon. Poytdk. 4: 178 

(1949). Type: Australia, Victoria, Barwon Heads, 1894, F. R. M. Wilson s.n. (H! (herb. Rasanen) - 

holotype; no medullary acids, usnic acid only). Plate 13, fig. 7. 

Thallus corticolous, pale green, caespitose, erect to subpendulous, up to 4 cm long, excep- 
tions to 6 cm; branching sparse, subdichotomous to irregular, additional side branchlets occur 
on narrow branches; branch width (0-5-)l-5(-10) mm, branches compressed, narrow and 
canaliculate or broad and flat, apices usually forked, narrow branches with fine pointed apices 
broad branches with dissected apices; surface shiny and smooth or matt and rugose, becoming 
coarse with chondroid strands showing at the surface, often horny in texture; holdfast delimited 
or diffuse (if growing in colonies); soralia absent. 

Apothecia common, laminal at the axil of bifurcating branches (to 5 mm diam.) or laminal 
towards the branch apices (0-5-1-0 mm diam.), marginal and subterminal on narrow canalicu- 
late branches (0-5-1-5 mm diam.), laminal on broad branches (2-10 mm diam.) at or near the 
apex margin, with smaller apothecia on the same branch near the centre; disc concave becoming 
plane, rarely convex; margin thick to thin, persistent, often inrolled; spores ellipsoid, straight or 
curved, 10-12(-16) x 4-5(-6) /xm. 

Chemistry. Sekikaic acid (± trace homosekikaic acid or trace divaricatic acid), or usnic acid 
only, or divaricatic acid and usnic acid. 

Remarks. The type material of R. glaucescens comprises a branch covered with tiny tufted 
specimens and a card with four specimens attached to it (which are here designated a, b, c, d). 
Two growth forms are represented on the card; a and d have bifurcate branches with small 
apothecia lying at the axil of the fork, and b and c have broader branches and bear terminal or 
subterminal apothecia, the latter with small subtending spurs. All of the thaUi tested contain 
sekikaic acid. The specimens are exceptionally small and are immature. 

The considerable polymorphism encountered in R. glaucescens presented much difficulty in 
interpretation, as there was no correlation with the chemical variation; a situation rather similar 
to that encountered by Kristinsson (1969) in dealing with Cetraria islandica in Iceland. Five 
different species names already existed for entities now included in the complex: R. glaucescens 
Krempelh., R. leiodea war . fastigiatula Miill. Arg., /?. brevis F. Wilson, R. lacerata Miill. Arg., 
and R. calicaris var. australica Rasanen. After the examination of large quantities of material 
belonging to this complex, both in the field and from herbarium collections, it became obvious 
that the four names additional to that of R. glaucescens (the earliest name) could not be 
maintained as species, because of the subtle intergradation between each of them (Plate 8, figs 
1-15). 

An examination of spores from all of the morphotypes failed to show any marked differences 
in spore size or shape. Each apothecium produced spores with considerable variation in both 



164 



G. NELL STEVENS 




RAMALINA IN AUSTRALIA 165 

characters, so that the ratio of breadth to length was never constant for any particular 
morphotype. The ratio of straight to bent spores was also variable with a tendency towards a 
greater proportion of bent spores in the coastal thalli. 

The taxonomic interpretation of this complexity of morphotypes offered two alternatives: (1) 
to regard each existing ecotype as a distinct species which would allow for the five existing 
species and would produce at least two other new taxa, or (2) to regard the whole group as one 
extremely variable species showing remarkable phenotypic plasticity in response to environ- 
mental pressures. The latter would incorporate numerous morphotypes with similar spores and 
anatomy, which occupied different habitats, yet showed intergradation between their extreme 
morphological forms (Plate 8, figs 1-15). The second alternative was chosen because variation in 
chemistry also occurs throughout the group, which made it impossible to satisfactorily find any 
point for delimitation based on the acids present. 

Consideration was given to applying varietal rank to the different ecotypes, but each ecotype 
would then have had a sekikaic acid race and an acid-deficient race. The division of R. 
glaucescens into two taxa based on presence or absence of medullary acids could be feasible, 
especially as thalli containing sekikaic acid occur mainly east of longitude 147°, whereas west of 
this latitude the majority of the specimens contain no acids. If a division of taxa was made this 
way then a mixed group of morphotypes would be placed in each class. The amount of sekikaic 
acid present in R. glaucescens specimens is very variable in quantity, and division of a species on 
so fine a difference as presence or absence of traces of sekikaic acid appeared unwarranted. 
Usually acid-deficient races are not distinguished taxonomically from their acid-producing 
partners. 

The extreme variability of this taxon (as described above) indicates it could be a rapidly 
expanding species in Australia. As it also occurs in New Zealand, investigation of material here 
may provide information to clarify how it should be divided, whether on its chemistry or its 
morphology. Until the matter is finally resolved this complex of morphotypes is regarded as a 
single, markedly polymorphic species. 

Two specimens collected from Walcha Road (31°S) contain divaricatic acid; and both sekikaic 
and divaricatic acids occur in several thalli collected at Nimmitabel (36°S); this anomaly cannot 
be explained. Both areas are at altitudes of over 1000 m and such habitats may produce 
physiological changes in the metabolism of R. glaucescens. No taxonomic rank was considered 
for the atypical divaricatic acid thalli. 

Distribution and habitat. Ramalina glaucescens has a wide distribution in the warm to cool 
temperate regions of eastern Australia (Fig. 13), occurring at altitudes from sea level to over 
1300 m. A large number of morphotypes are found as illustrated in Plate 8, figs 1-15. A general 
intergradation of forms can be seen which do not form a continuum but merge from several 
directions. 

Thalli with a morphology which is larger, but otherwise similar to that of the type of R. 
glaucescens, are found along the inland mountains west of the eastern escarpment in New South 
Wales and in the mountainous area of central Victoria, as well as in the central and eastern 
mountains of Tasmania (Plate 8, figs 2, 3, 4, 5). All of the specimens examined contain sekikaic 
acid in varying amounts, except for a few from western Victoria and South Australia which are 
acid-deficient. 

Intergrading with this morphotype is a narrower, sometimes canaliculate morph which 
often has subsidiary branchlets extending from the main branches; it has both terminal and sub- 
terminal apothecia. The distribution of this ecotype is in the Eastern Highlands (with a disjunct 
appearance in the Grampians - type location of R. leiodea var. fastigiatula) and in the highlands 
of Tasmania (Plate 8, figs 5, 6, 7, 9, 13, 14). In its narrowest form the branches become subterete 
and very fine, bearing small apothecia; the apices of the branches are sharply pointed (Plate 8, 
figs 11, 12); this spiky form occurs in mountain habitats above 1400 m elevation in the temperate 
region, e.g. it occurs at Mt Aggie and the Brindabella Ranges in New South Wales and at Mt 
Wombargo in Victoria, and at lower altitudes at Buchan Caves, Victoria, and in Tasmania. All 
collections were from twigs of Epacridaceae bushes, so it is not clear whether low temperatures 



166 



G. NELL STEVENS 




Fig. 13 Distribution of JR. glaucescens: • sekikaic acid race; O acid-deficient race; * divaricatic acid race. 

or a particular bark substrate has an effect on this phenotype's distribution. Sekikaic acid was 
always present in such thalli. 

One ecotype of R. glaucescens grows along the southern Victorian coastUne; it has a short 
broad thallus with large terminal or laminal apothecia and is acid-deficient. A few specimens 
resembling this morphology have been collected from inland in eastern Australia, but that 
material contains sekikaic acid and is not so coarsely textured as the coastal thalli. This ecotype 
resembles the type of R. brevis (Plate 8, fig. 15). 

Another acid-deficient ecotype occurs inland at low altitudes in Victoria and South Australia, 
but the thallus is generally narrower than that of the coastal ecotype and the apothecia are not as 
large; however, intergradation between the two is evident at some locations (Plate 8, fig. 8). The 
presence of small branchlets on the main branches resembles the eastern highlands ecotype. The 
type of R. calicaris var. australica belongs to this ecotype. In some habitats the thallus becomes 
quite coarse in texture with chondroid strands showing at the surface; such characteristics are 
apparent in the material from the arid zone in Western Australia, including the type collection of 
R. lacerata. 

The largest morphotype in the R. glaucescens complex is found in western Victoria and 
eastern South Australia, with a few specimens collected in western New South Wales. The latter 
specimens contain sekikaic acid but all other material proved to be acid-deficient. Most of the 
ecotypes of R. glaucescens have been found to intergrade with this large morphotype (Plate 8, 
fig. 1). It is normally sterile although pycnidia have been detected in some of these specimens. 

Four of the type specimens here included in R. glaucescens were initially collected within 100 



RAMALINA IN AUSTRALIA 167 

km radius of one another in Victoria, which illustrates the ecophenotypic response to different 
habitats. Table 9 summarises the morphological variation that is found in the different ecotypes. 
This taxon is entirely corticolous, the most common phorophytes being Acacia spp. and 
Bursaria spinosa. It occurs as dense colonies along the branches of isolated trees in pastureland 
or in open-forest communities. In some localities the entire population may be small and 
immature (as found in the type material of both R. glaucescens and R. leiodea var. fastigiatula) , 
whereas in other areas only large mature specimens occur. 

World distribution. Australia and New Zealand. 

New South Wales: Ben Lomond, 30°00'S, 151°42'E, on Acacia sp., 1980, GNS (BRIU3249NS); Sherwood 
Ck, 30°03'S, 153°03'E, on lemon tree, 1978, HS (CBG7905095); Walcha, 10 km W of, 30°58'S, 151°30'E, 
on Acacia sp. , 1982, GNS (BRIU4100NS); Abercrombie Caves, 33°32'S, 149°15'S, on Leptospermum sp. , 
1978, JAE (JAE 4640); Rockley, 33°42'S, 149°34'E, E. Boorman (G); Black Springs 10 km N of, 33°49'S, 
149°48'E, on Acacia sp. , 1984, NCS (BRIU4292NS); Black Springs 5 km S of, 33°52'S, 149°48'E, on Acacia 
sp., 1984, NCS (BRIU4283NS); Abercrombie R. 34°12'S, 149°45'E, on Pinm sp., 1984, NCS 
(BRIU4284NS); Wombeyan Caves, 34°18'S, 150°02'E, E. Cheel (NSW L4294); Taralga, 37 km N of, 
34°24'S, 149°49'E, on fence posts, 1977, JAE (JAE 3165); Crookwell, 34°28'S, 149°30'E, on Acacia sp., 
1980, GNS (BRIU3489NS); Yass, 26 km S of, 35°00'S, 148°50'E, on Casuarina sp., 1975, HS 
(CBG058278); Little Peppercorn Ck, 35°35'S, 148°37'E, on Bursaria spinosa, 1979, HS (CBG7906928); 
Rules Point, 35°43'S, 148°31'E, on shrub, 1979, HS (CBG7907045); Lake Eucumbene, 36°10'S, 148°50'E, 
on Hymenanthera sp., 1971, L. Craven (MEL1020182); Adaminaby, 36°12'S, 148°50'E, on Briar, 1984, 
NCS (BRIU4287NS); Countegany, 36°12'S, 149°29'E, on Acacia melanoxylon, 1976, JAE (JAE 1925); 
Cooma 46 km S of, 36°14'S, 149°05'E, on Acacia sp., 1978, JAE (JAE 5440); Nimmitabel, 36°31'S, 
149°15'E, on Prunus sp., 1980, GNS (BRIU3215NS); Holts Flat, 36°40'S, 149°15'E, on Acacia sp., 1980, 
GN5 (BRIU3310NS); Delegate, 37°03'S, 148°57'E, on Acacia sp., 1980, GNS (BRIU3490NS). Australian 
Capital Territory: Ginninderra Ck, 35°05'S, 149°03'E, 1977, HS (CBG066912); Uriarra Crossing, 35°08'S, 
148°57'E, on Casuarina sp., 1974, JAE (JAE 628); Blundells Ck Rd., 35°21'S, 148°50'E, on Acacia sp., 
1977, D. Verdon (CBG8008538); Mt Aggie, 35°27'S, 148°46'E, on Hymenanthera dentata, 1979, JAE 
(JAE 5870). Victoria: Walwa, 9 km S of, 35°55'S, 147°45'E, on Acacia sp. , 1984, NCS (BRIU4338NS); SW 
of Corryong, 36°16'S, 146°50'E, on Briar, 1984, NCS (BRIU4288NS); Myrtleford, 19 km S of, 36°39'S, 
146°42'E, on Acacia sp., 1984, NCS (BRIU4290NS); Tawonga, 36°41'S, 147°08'E, on Prunus sp., 1984, 
NCS (BRIU4293NS); Bright, 36°44'S, 146°58'E, on copper beech, 1984, NCS (BRIU4286NS); Mt 
Wombergo, 36°36'S, 148°11'S, on Hymenanthera sp., 1966, RF. (MEL1026138); Bonang, 37°12'S, 
148°43'E, on Malus, 1980, GNS (BRIU3491NS); Kyneton, 37°15'S, 144°28'E, 1897, F. R. M. Wilson, 
(MEL 9440); Newlyn, 37°25'S, 144°00'E, on Crataegus sp., 1981, RWR (BRIU2334RR); Buchan Caves, 
37°30'S, 148°10'E, 1965, RF (MEL1026151); Omeo, 7 km N of, 37°04'S, 147°38'E, on Acacia sp., 1984, 
NCS (BRIU4285NS); Skipton, 11 km S of, 37°47'S, 143°22'E, on Hymenanthera sp., 1984, NCS 
(BRIU4291NS); Buninyong, 37°42'S, 143°40'E, on trees, F. R. M. Wilson (MEL 9479); Steiglitz, 
Moorabool R dam, 37°53'S, 144°11'E, on twigs, 1968, GB (H032786); Yallourn, Storage dam,38°10'S, 
146°20'E, 1975, 7?F(MEL1013745); Koweerup, 12 km NE of, 38°10'S, 145°36'E, on Pinus sp., 1984, NCS 
(BRIU4336NS); Loy Yang, 38°11'S, 146°37'E, on Acacia sp., 1984, NCS (BRIU4337NS); Billywing 
Forest, 37°10'S, 142°10'E, Acacia baileyi, 1981, RWR (BRIU2429RR); Dartmoor, Crawford Lake, 
37°56'S, 141°25'E, 1978, T. Muir (MEL1025115); Macarthur, 16 km W of, 37°58'S, 141°48'E, on 
Leptospermum sp., 1969, A. Orchard (SA97647210); Sale, 38°07'S, 147°04'E, on Acacia sp., 1980, GNS 
(BRIU3208NS); Tyrendarra, 38°15'S, 141°50'E, on tree, 1951, A. C. Beauglehole (MEL 9487); Lake 
Beeac, 38°12'S, 143°38'E, on Briar, 1984, NCS (BRIU4340NS); Barwon Hds, 38°20'S, 144°30'E, 1894, F. 
R. M. Wilson (H); Gunnamatta Beach, 38°20'S, 144°45'E, on Pimelea sp., 1983, GNS (BRIU3680NS); 
Strzelecki South, 38°20'S, 145°58'E, on dead wood, 1971, Gfi <fe/C(H032630); Westernport Bay, 38°22'S, 
145°20'E, on Avicennia marina, 1978, GNS (BRIU4250NS); Warrnambool, 38°23'S, 142°31'E, on 
Bursaria spinosa, 1887, F. R. M. Wilson (NSW L4091); Pirron Yalook, 38°21'S, 143°26'E, on shrub, 1952, 
A. C. Beauglehole (MEL1023376); Yarram, 17 km NNW of, 38°24'S, 146°38'E, on Acacia sp., 1980, HS 
(CBG8002966); Korumburra, 38°26'S, 145°49'E, on Acacia sp., 1980, GNS (BRIU3301NS); Forrest, 
upper Barwon R, 38°32'S, 143°45'E, on 'prickly moses' bush, 1952, A. C. Beauglehole (MEL1023362); 
Gellibrand R. falls, 38°40'S, 143°10'E, on tree, 1952, A. C. Beauglehole (MEL1023385); Skenes Ck, 
38°43'S, 143°45'E, on Helicrysum dendroideum, 1983, M. Seaward 104633; Wilsons Promontary, 39°06'S, 
146°12'E, on Acacia sp., 1980, GNS (BRIU3207NS). South Australia: Yalata Roadhouse, 31°39'S, 
129°03'E, on bushes, 1980, AA (BRIU4341NS); Swan Reach, 24 km S of, 34°37'S, 139°30'E, 1971, K. 
Czornij (SA97414420); Springton, 6 km W of, 34°39'S, 139°06'E, on Hakea sp., 1975, JAE (JAE 884); 



168 



G. NELL STEVENS 



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RAMALINA IN AUSTRALIA 169 

Curramulka 5 km W of, 34°43'S, 137°45'E, on Casuarina sp., 1977, JAE (JAE 3731); Kuitpo Forest, 
35°14'S, 138°36'E, windfall, 1968, RWR (BRIU3032RR); Myponga, 35°24'S, 138°28'E, 1968, V. Cruick- 
shank (SA97414419); Clarendon, 35°06'S, 138°40'E, 1968, V. Cruickshank (SA97418422); Strathalbyn, 
35°20'S, 138°50'E, 1968, V. Cruickshank (SA97414424); Salt Ck, 35°28'S, 139°09'E, on dead gum, 1967, 
GB (H032783); Deep Ck, 35°38'S, 138°12'E, on Casuarina sp., 1979, RV/R (BRIU1863RR); Messent 
N.P., 35°28'S, 139°10'E, 1969, J. Carrick (SA97416171); Hindmarsh Falls, 35°29'S, 138°38'E, 1968, V. 
Cruickshank (SA97416168); Kangaroo I. Lockwood Cnr. 35°39'S, 137°38'E, on tree, 1967, G. Jackson 
(SA97647604); The Gap, 36°41'S, 140°40'E, 1973, M. Beck (H032671); Kingston, 60 km N of, 36°50'S, 
139°51'E, P. Copley (SA97648420); Milbrook Reservoir, 34°50'S, 138°47'E, on Hakea sp., 1969, RWR 
(BRIU1780RR); Penola, 37°23'S, 140°49'E, on twigs, 1971, RWR (BRIU2035RR). Western Australia: 
Eucla, 31°43'S, 128°55'E, /. Oliver (G, M). Tasmania: Trowutta, 41°02'S, 145°05'E, on Malm sp., 1981, 
GK 384/81; Beaconfield, 41°12'S, 146°45'E, on Acacia dealbata, 1980, GK 214/80; Prossers Forest Rd, 
4ri5'S, 147°18'E, on Malus sp., 1966, GB (H032777); Avoca, Lochaver, 41°41'S, 147°50'E, on Prunus 
sp., 1978, GK 1/78; Cressy, 41°43'S, 147°02'E, on dead Acacia, 1964, JC (H032576); Lake Augusta, 
41°50'S, 146°34'E, on twigs, 1973, GB (H032591); Great Lake Plateau, 41°57'S, 146°40'E, on bush, 1969, 
GB (H032628); Macquarie R, 42°08'S, 147°33'E, on small bushes, 1966, GB&JC (H032780); Lake Tooms 
Rd, 42°20'S, 147°28'E, on small bushes, 1974, GB & Gilbert, (H032791); Bothwell, 42°23'S, 147°00'E, 
1968, GB & JC (H032593); Spring Hill, 42°24'S, 147°15'E, 1963, GB (H032770); Clyde Valley, 42°27'S, 
146°52'E, on Acacia sp., 1975, K. Davies (H032629); Ouse, 42°28'S, 146°44'E, on Malus sp., 1980, GNS 
(BRIU3340NS); Bushy Park, 42°43'S, 146°53'E, on Exocarpus cupressiformis , 1980, G/C 31/80. 



8. Ramalina /itorea N. Stevens Plate 7, fig. 8. 

in Lichenologist 18: 185 (1986). Type: Australia, Queensland, Mackay, Blacks Beach, 1979, Stevens 
3129NS (MEL! - holotype; evernic acid, lecanoric acid and usnic acid). 

Thallus saxicolous, pale yellow-green, small, caespitose, rigid, 0-5-2-0 cm high; branching 
subdichotomous to irregular, small branchlets or spinules arise laterally from the main branches; 
branch width 0-2-1 -0 mm, subterete to terete or rarely flattened, narrow, apices pointed, 
fragile; surface shiny to matt, without pseudocyphellae, smooth; holdfast diffuse; soralia absent. 

Apothecia common, marginal or lateral below the apices, disc 1-0-2-0 mm diam., plane; 
margin thin to very thick; spores small, oval or ellipsoid, straight, 8-12 x 4-4-5 fxm. 

Chemistry. Evernic acid (M), lecanoric acid (t), ± sekikaic acid (t), and usnic acid. 

Remarks. Some specimens of R. litorea resemble the Hawaiian species R. microspora Krem- 
pelh. However, the latter species contains divaricatic acid, whereas R. litorea contains evernic 
acid. Herbarium material labelled R. microspora from Mauritius and Rodrigues (in the Indian 
Ocean, 20°S) containing evernic acid and morphologically somewhat similar to the Australian 
taxon have been treated as R. litorea by the author. Further investigation may prove that R. 
microspora comprises two chemical races disjunctly distributed in the Pacific and Indian 
Oceans. 

Distribution and habitat. The full distribution range of R. litorea (Fig. 14) is unknown as only 
five collections have been made in an area from latitudes 14°40'-24°20'S. This saxicolous species 
has been collected from coastal cliffs, composed of a variety of rock types (granite, siltstone, 
slates, rhyohtic, breccia, and tuff) indicating it is not substrate specific. Only small colonies grow 
on the cliff faces, usually at 10-20 m above beach level; although on Lizard Island (14°S, the 
most northern collection) a specimen was collected at 360 m elevation. 

World distribution. Mauritius, Rodrigues Island, and Australia. 

Queensland: Lizard I. 14°40'S, 145°28'E, on granite (360 m elev.), 1974, R. L. Specht (BRIU3780NS); 
Cape Hillsborough, 20°56'S, 149°03'E, on rhyolitic breccia (10 m elev.), 1983, GNS (BRIU4124NS); 
Mackay, Blacks Beach, 21°04'S, 149°12'E, on siltstone (10 m elev.), 1979, GNS (BRIU3129NS); Great 
Keppel \, 23°10'S, 150°58'E, substrate unknown, 1976, U. Allen (H032240); Double Head, 23°08'S, 
150°47'E, on slates (15 m elev.), 1984, NCS (BRIU4307NS); Emu Park, 23°15'S, 150°30'E, on slates (10 m 
elev.), 1984, NCS (BRIU4308NS); Agnes Water, 24°20'S, 151°58'E, on rhyolitic agglomerate (5 m elev.) 
1984, NCS (BRIU4277NS). 



170 



G. NELL STEVENS 




Fig. 14 Distribution ofR. litorea. 



9. Ramalina peruviana Ach. Plate 9, figs 1-2. 

Lichenogr. Univ. : 599 (1810). Type: South America, Peru, Lagasta s.n. (H-ACH! -holotype; BM!, UPS! 

- isotypes; sekikaic acid aggregate, and usnic acid). 
Ramalirta farinacea var. dendroides Miill. Arg. in Flora, Jena 66: 21 (1883). Type: Australia, New South 

Wales, Ash Island (Hunter River), 1883, Ford s.n. (G! - holotype; sekikaic acid aggregate and usnic 

acid). Plate 14, fig. 2. 
Ramalina farinacea var. squarrosa Miill. Arg. in Flora, Jena 66: 21 (1883). Type: Australia, Queensland, 

Rockhampton, 1883, Thozets.n. (G!- holotype; sekikaic acid aggregate and usnic acid). Plate 14, fig. 1. 

Thallus corticolous, pale green to grey-green, tufty, resilient, erect to subpendulous, up to 6-0 
cm long; branching subdichotomous to irregular, often intricate with the production of dense 
fragile branchlets; branch width up to 1-0 mm, branches flat, compressed becoming angularly 
subterete to terete distally, branches often slightly twisted, apices sharp to blunt, often broken; 
surface matt, rarely shiny, smooth to rugose, weakly pseudocyphellate either basally or along 
the entire length; holdfast delimited or diffuse; soralia punctiform, numerous, marginal or 
lateral, sometimes apical (Plate 1, fig. 2), mounds of soredia often produce small fibrils (Plate 3, 
fig. 7). 

Plate 9 Thallus growth forms in the Australian Ramalinae. Scale in mm. Fig. 1 R. peruviana (dense 
form). Fig. 2 R. peruviana (open form). Fig. 3 R. reducta (lower surface). Fig. 4 R. tenella. Fig. 5 R. 
unilateralis (palmate form). Fig. 6 R. unilateralis (large, intricate form). Fig. 7 R. whinrayi. Fig. 8 R. 
celastri subsp. celastri (narrow form). 



RAMALINA IN AUSTRALIA 171 

iiMiiinilititlimlmiliiiil!iiilniiliiiiHiiiliiiiliiiiliiiiliiiimi 




172 G. NELL STEVENS 

Apothecia rare, marginal, lateral, disc 2-0 mm diam., concave, margin thick, often crenate; 
spores narrow, fusiform, straight or rarely curved, 14-16(-18) x 3-5-4-5 /xm. 

Chemistry. Homosekikaic acid (M), sekikaic acid (M), ramalinolic acid (m/t), 4'-0- 
demethylsekikaic acid (m/t), 4'-0-methylnorsekikaic acid (m/t), and usnic acid. 

Remarks. The holotype material from H and the isotypes material held at BM and UPS are all 
small pieces and the morphological characters are not well defined, faint pseudocyphellae occur 
on the basal portion of the branches, and the soralia are not very clear. The Australian material 
does not closely resemble the morphology of the type material but Krog & Swinscow (1976) 
commented that some of the East African specimens appeared to differ from the type specimen. 
When comparing East African material with Australian no differences could be found. 
Ramalina peruviana material examined from Uruguay and Brazil closely resembled Australian 
specimens and also displayed considerable variation in morphology. 

Two previously accepted infra-specific taxa from Australia are reduced to synonymy with R. 
peruviana; these are R. farinacea var. dendroides and R. farinacea var. squarrosa, both of which 
are regarded as ecophenotypes of R. peruviana. 

Ramalina tenella is the only species in Australia which may be mistaken for R. peruviana, but 
the acid difference sets them apart, the former containing salazinic acid. The Japanese taxon R. 
intermediella Vainio closely resembles the morphology of R. peruviana and contains similar 
acids, but it does not occur in Australia. 

Distribution and habitat. Ramalina peruviana is mainly subtropical to warm temperate in 
distribution (Fig. 15). North of the Tropic of Capricorn it occurs inland at 1000 m elevation at 
both Atherton Tableland (17°S) and Undara Crater (19°S). Such elevations in the tropics 
correspond with the reported distribution of the species in the West Indies (17°N) at altitudes 
between 600-1000 m (Landron, 1972), and in East Africa (10°N-10°S) between 1000-1800 m 
(Krog & Swinscow, 1976) allowing for latitudinal adjustment. 

Four ecotypes can be distinguished in the Australian taxa although intergradation occurs 
between each form: (i) the coastal form (which includes R. farinacea var. squarrosa type 
material), (ii) a montane form, (iii) a densely branched form found in open-forest habitats in 
Queensland and (iv) an open sparsely branched form in the open-forest habitats of south-east 
Queensland and New South Wales (which includes R. farinacea var. dendroides type material). 
The differences are shown in Table 10. 

Specimens of these four ecotypes were chemically analysed to assess if there was any 
correlation between their chemistry and the morphological differences. None was found, but 
variation in the quantity of some of the minor metabolites was obvious. This variation could be 
correlated with inland and coastal locations, i.e. coastal specimens had only trace amounts of 
4'-0-methylnorsekikaic acid, but specimens from inland sites contained noticeably more. 

J0rgensen (1977) commented that R. peruviana may need to be subdivided because of the 
number of morphotypes present, but from the survey of Australian material it is felt that any 
subdivision is unwarranted because of the intergradation between ecotypes. 

All collections in Australia have been corticolous. The main phorophytes for the coastal 
ecotype are the mangroves Ceriops tagal and Rhizophora stylosa. Once these species reach their 
distribution limit at 28°S, R. peruviana is found on various coastal trees. In the mountains this 
taxon occurs on several rain-forest tree species and in the open-forest habitat, shrubs such as 
Lantana are the host. This taxon is not substrate specific; in South America it has been found on 
fence posts (Osorio, in Utt.). 

World distribution. Africa, Australia, New Zealand, several Pacific Islands, South America, 
and Tristan da Cunha. 

Queensland: Atherton, 5 km N of, 17°14'S, 145°29'E, 1970, E. Dahl (0); Atherton, 2 km SE of, 17°16'S, 
145°30'E, on Casuarina sp., 1983, HS (CBG8302451); Undara Crater, 18°23'S, 144°43'E, on rain-forest 
tree, 1977, NCS (BRIU3546NS); Forty Mile Scrub, 18°03'S, 144°53'E, on rain-forest tree, 1983, GNS 
(BRIU4115NS); St Lawrence, 22°20'S, 149°32'E, on Ceriops tagal, 1978, NCS (BRIU2678NS); Glad- 
stone, South Trees Inlet, 23°52'S, 151°19'E on Ceriops tagal, 1980, GNS (BRIU3592NS); Tannum Sands, 



RAMALINA IN AUSTRALIA 

iTo 



173 




Fig. 15 Distribution of R. peruviana. 



23°58'S, 151°23'E, on Rhizophora stylosa, 1975, RWR (BRIU876RR); Turkey, 24°06'S, 151°38'E, on 
Ceriops tagal, 1975, GNS (BRIU1154NS): Jimna, 26°40'S, 152°28'E, on Araucaria cunninghamii, 1980, 
RWR (BRIU2197RR); Maleny, 26°46'S, 152°51'E, on dead tree, 1982, GNS (BRIU3708NS); Bunya Mts, 
26°50'S, 151°40'E, on shrubs, 1979, GNS (BRIU2971NS); Mt Mee, 27°05'S, 152°42'E, on Flindersia sp., 
1979, GNS (BRIU3324NS); Mt Byron, W of 27°06'S, 152°37'E, on Araucaria cunninghamii 1982, GNS 
(BRIU3624NS); Brisbane, Serpentine Ck, 27°22'S, 153°08'E, on Ceriops tagal, 1975, GNS 
(BRIU1097NS); Stradbroke I. Amity Pt, 27°24'S, 153°25'E, on Lumnitzera racemosa, 1977, GNS 
(BRIU2050NS); Brookfield, 27°30'S, 152°55'E, on Citriobatus pauciflora, 1983, GNS (BRIU3694NS); St 
Lucia, 27°30'S, 153°00'E, on Bauhinia alba, 1976, GNS (BRIU3322NS); Worlds End Pocket, 27°31'S, 
152°45'E, on Mallotus celaoxy hides, 1983, P. Forster (BRIU3705NS); Pt Talburpin, 27°39'S, 153°18'E, on 
Avicennia marina, 1980, GNS (BRIU3548NS); Little Rocky Pt, 27°43'S, 153°21'E, on Ceriops tagal, 1977, 
GNS (BRIU480NS); Stradbroke L south point, 27°44'S, 153°25'E, on mangrove, 1975, H. T. Clifford 
(BRIU3074RR); Cunninghams Gap, 28°03'S, 152°24'E, on rain-forest tree, 1975, NCS (BRIU3695NS); 
Binna Burra, 28°12'S, 153°11'E, on rain-forest tree, 1976, GNS (BRIU3693NS); O'Reillys, 10 km N of, 
28°13'S, 153°06'E, on Gympie tree, 1983, GNS (BRIU3724NS). New South Wales: Wilsons Ck, 28°34'S, 
153°27'E, on rain-forest tree, 1980, GNS (BRIU3320NS); Goonengarry, 28°37'S, 153°28'E, on Citriobatus 
pauciflora, 1980, GNS (BRIU3319NS); The Channons, 28°39'S, 153°16'E, on Lantana, 1982, GNS 
(BRIU3526NS);Lismore,28°48'S, 153°16'E,on^cfldflsp., 1976,/?. Moorf/e (BRIU2855NS);Nambucca, 
12 km SSW. of, 30°41'S, 153°00'E, on Avicennia marina, 1982, GNS (BRIU3670NS); Taree, Saltwater, 
31°54'S, 152°34'E, on tree, 1977, JAE (JAE 3999); Hunter R. Ash L 32°56'S, 15r46'E, 1883, E. Forde 
(G); Broulee, 35°52'S, 150°09'E, on shrubs, 1970, JAE (JAE 128); Wagonga Inlet, 36°12'S, 150°08'E, 
1908, G. H. Halligan NSW L4304). 



174 



G. NELL STEVENS 









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RAMALINA IN AUSTRALIA 175 

10. Ramalina reducta Krog & Swinscow Plate 9, fig. 3. 

in Norw. J. Bot. 23: 172 (1976). Type: Ethiopia, Bale Province, Dinshu, Tapper933 (BM! -holotype; O! - 
isotype; psoromic acid and usnic acid). 

Thallus corticolous, grey-green or pale green, erect, flaccid, up to 4 cm long; branching 
sparse, initially a palmate lobe producing several multi-divided lacinae; branch width 10 mm at 
base, separate lacinae to 5 mm, branches compressed, flat, apices tattered or capillaceous; 
surface matt, rugose, often cracking; a continuous lower cortex lacking with the medulla 
overlaid by cartilaginous strands forming a reticular pattern, some strands becoming detached at 
one end; holdfast delimited; soredia pale green, produced from the exposed medulla. 

Apothecia rare, marginal, disc to 2-5 mm diam, concave to plane, margin entire; spores few, 
ellipsoid, curved, 13-18 x 4-5(-6) /xm. 

Chemistry. Psoromic acid and usnic acid. 

Remarks. The Australian specimens are much larger than those examined from East and South 
Africa and cannot be regarded as 'inconspicuous' as described by Krog & Swinscow (1976). This 
species is easily recognised by the anastomosing strands of cartilaginous hyphae on the exposed 
lower surface. The production of psoromic acid distinguishes it from any other Ramalinae in 
Austraha. 

Distribution and habitat. This taxon has a restricted distribution range (Fig. 16) . It occurs in the 
open-forest uplands of the New England Plateau, a granitic/basaltic tableland rising to over 





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176 G. NELL STEVENS 

1300 m in the subtropics. The narrow ranges of latitude and longitude in R. reducta distribution 
coincide with a 12°C mean maximum annual temperature isotherm. The distribution area 
experiences wet summers and drier winters, indicating that the summer may be the growing 
period for this species. This particular set of environmental factors is not found in other areas. 

To the north only individual peaks reach an elevation of over 1300 m and these are covered in 
rain-forest; and to the south, where the Australian Alps attain a height of over 2000 m, the 
rainfall is either uniform throughout the year or occurs during the winter. Seasonality may be a 
factor which inhibits the occurrence of R. reducta further south. 

Krog & Swinscow (1976) recorded East African material from the upper montane forests 
between 2600 m and 3100 m altitude, which when adjusted for latitude is comparable with the 
occurrence of R. reducta in Australia. 

Ramalina reducta was collected horn Acacia, Casuarina, and Banksia spp. growing in well lit 
open-forest communities. A large monospecific community of this taxon was found at Ebor 
Falls, New South Wales, growing on all three phorophytes, in an area exposed to spray from the 
falls under certain wind conditions. In this environment thalU reached 4 cm in length and many 
produced apothecia, indicating that such a habitat promoted optimal growth. No fertile thalli 
were reported from the East African material. 

World distribution. Africa and Australia. 

New South Wales: Glen Innes, 12 km E of, 29°42'S, 151°50'E, on Acacia parvipinula, 1983, GNS 
(BRIU3782NS); Ben Lomond, 30°00'S, 151°42'E, on dead Acacia sp., 1980, GNS (BRIU3237NS); 1976, 
JAE (JAE 2435); Coutts Water, Ebor-Dorrigo Rd, 30°22'S, 152°30'E, on shrubs, 1982, GNS 
(BRIU3633NS); Ebor Falls, 30°25'S, 152°20'E, on Banksiasp., 1982, GN5 (BRIU3632NS); on cicada sp., 
1982, GNS (BRIU3698NS); on Casuarina sp., 1982, GNS (BRIU3699NS); Walcha Road, 30°59'S, 
151°30'E, on Acacia sp., 1982, GNS (BRIU3700NS); Topdale, 25 km E of Nundle, 31°30'S, 151°25'E, on 
Acacia sp., 1984, NCS (BRIU4297NS); Glenbawn Dam, 32°05'S, 151°00'E, on Acacia sp., 1981, AA 
(BRIU3611NS). 

11. Ramalina tenella Miill. Arg. Plate 9, fig. 4 

in Flora, Jena 62: 162 (1879). Type: Brazil, Sao Paulo, Apiahy, 1877, Puiggari 152 (G! - holotype; salazinic 
acid, protocetraric acid (trace), three unknowns, usnic acid). 

Thallus corticolous, pale green to yellow-green, erect, rigid, dehcate, up to 3-0 cm high; 
branching subdichotomous with short lateral branches sometimes producing numerous distal 
branchlets; branch width to 1-0 mm, flat basally thereafter subterete; surface matt; short, hnear 
pseudocyphellae; holdfast delimited; soralia apical and lateral on main branches and apical on 
secondary branches. 

Apothecia unknown. 

Chemistry. Salazinic acid ± protocetraric acid, three unknowns, and usnic acid. 

Remarks. This taxon has a distinctive growth form when the branching is sparse; when densely 
branched it resembles R. peruviana. The presence of salazinic acid sets it apart from the sekikaic 
acid taxon R. peruviana. Krog & Swinscow (1976) stated that R. tenella resembled R. 
dendriscoides Nyl. , a species which does not occur in Australia, although some early collections 
of R. peruviana were given the name R. dendriscoides (Appendix 1). 

Distribution and habitat. Ramalina tenella occurs in isolated, high rainfall pockets along the 
Queensland coast between 16°S and 24°S (Fig. 17). Uncommon in occurrence, it grows profusely 
in the areas it inhabits. Material collected from the more northern latitudes always contained 
three unknown compounds which did not occur in tl;e material from latitudes 2rS-24°S. Lack 
of bulk material of this species prevented the identification of the unknowns by chemical 
analysis. Specimens lacking the unknowns were usually found to produce lateral soralia in 
addition to apical soralia, in contrast to the more northern collections which rarely produced 
lateral soralia. 

Ramalina tenella grows on the twigs of Ceriops tagal and Rhizophora stylosa in sheltered but 
well-lit habitats within the mangrove community. It is found in association with R. subfraxinea 



RAMALINA IN AUSTRALIA 



177 





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var. confirmata, R. pacifica, R. exiguella, R. nervulosa var. nervulosa, and /?. nervulosa var. 

World distribution. Africa, Australia, New Hebrides, and South America. 

Queensland: Daintree R, 16°15'S, 145°21'E, on Ceriops tagal, 1975, D. Tarte (BRIU2253NS); Mossman, 
Saltwater Ck, 16°25'S, 145°25'E, on Ceriops tagal, 1983, C. McCracken (BRIU4300NS); Mossman, Rifle 
Club, 16°28'S, 145°25'E, on Ceriops tagal, 1983, GN5(BRIU4127NS); Cairns, 16°45'S, 145°45'E, 1974, D. 
Tarte (BRIU2470NS); Murray R, 18°05', 146°01'E, on Ceriops tagal, 1978, D. Tarte (BRIU3122NS); 
Hinchinbrook I, 18°20'S, 146°15'E, on Ceriops tagal, 1979, GNS (BRIU3134NS); Eimeo, 21°09'S, 
149°10'E, on Ceriops tagal, 1977, GNS (BRIU2791NS); on Rhizophora stylosa, 1976, GNS 
(BRIU2469NS); Corio Bay, 23°00'S, 150°45'E, on Ceriops tagal, 1975, D. Tarte ^BRIU1172NS); 
Gladstone, Wiggins Ck, 23°50'S, 151°15'E, on Ceriops tagal, 1980, GNS (BRIU3549NS); Turkey, 24°06'S, 
151°38'E, on Rhizophora stylosa, 1975, GNS (BRIU1282NS). 



12. Ramalina unilateralis F. Wilson Plate 9, figs 5-6. 

in Victorian Nat. 6: 69 (1889). Type: Australia, Victoria, Warrnambool, F. R. M. Wilson 432 (G! - 
lectotype herein designated; divaricatic acid and usnic acid). 

Thallus corticolous, pale green, caespitose to subpendulous, up to 4 cm long, exceptionally to 
7 cm; branching irregular, dense; branch width 1-3 mm, flat to subterete, sometimes palmate at 
the base to 8 mm broad, distally producing numerous fine lacinae; surface cartilaginous, smooth 
and shiny, rarely matt; holdfast delimited; soralia occur in eroded patches at intervals along the 



178 G. NELL STEVENS 

lower surface where the branches have split and flattened out (Plate 1, fig. 6) and also at the 
apices. 

Apothecia rare, disc 2-0 mm diam., concave to plane; margin entire; spores ellipsoidal, 
straight or rarely curved, 10-12 x 4-4-5 fxm. 

Chemistry. Divaricatic acid and usnic acid. 

Remarks. The specimens held at G are regarded as the lectotype of R. unilateralis as no other 
Wilson material appears to exist. It is thought that the type may have been lost with numerous 
other Wilson types in 1907 (Filson, 1976). 

The morphology of some R. pollinaria specimens resembles the morphology of this taxon, and 
it was this name that was given to R. unilateralis specimens by early lichenologists. However, R. 
pollinaria contains evernic acid, whereas R. unilateralis contains divaricatic acid. 

Some specimens oiR. canariensis resemble R. unilateralis; however, the texture of the surface 
is different. Reduced growth forms of the latter taxon may be mistaken for R. fimbriata, but this 
species is saxicolous in Australia whereas R. unilateralis is corticolous and generally much larger 
in size. All three species contain divaricatic acid. 

Distribution and habitat. Collections of this cool temperate species in Australia have been made 
from sea-level to elevations of up to 1150 m (Fig. 18). Although corticolous in Australia R. 
unilateralis becomes saxicolous at colder latitudes, e.g. Macquarie I. (55°S) and Tierra del Fuego 
(54°S). It appears to tolerate a broad range of environments. The type material was collected at 




Fig. 18 Distribution of /?. unilateralis. 



RAMALINA IN AUSTRALIA 179 

sea-level, and other coastal sites are Lakes Entrance, Westernport Bay, and Portland; inland 
collections above 1000 m were at Nimmitabel, New South Wales, and Old Man's Head, 
Tasmania. Material cited from South Africa was collected at elevations of 2000 m at latitude 
34°S, and from Chile, latitude 53°30'S at an elevation of 800 m. 

World distribution. South Africa, Australia, and South America. 

New South Wales: Nimmitabel, 36°31'S, 149°17'E, on Banksia sp., 1976, JAE (JAE 1623); Nimmitabel, 
36°31'S, 149°15'E, on Prunus. 1980, GNS (BRIU3781NS). Australian Capital Territory: Mt Coree, 
35°20'S, 148°50'E, on tree, 1970, E. Dahl{0). Victoria: Bonang, 17 km N of, 37°03'S, 148°49'E, on Acacia 
sp., 1978, D. Verdon (CBG7811003); Billywing Forest, 37°10'S, 142°30'E, on Acacia baileyi, 1981, RWR 
(BRIU2451RR); Spring Ck Gap, 37°15'S, 148°30'E, on Acacia sp., 1978, JAE (JAE 5137); Metung, 
37°53'S, 147°51'E, on tree, 1889, F. R. M. Wilson (MEL 9420); Upper Maffra, 37°54'S, 146°51'E, on 
conglomerate, 1886, F. R. M. Wilson (NSW L4312); Hastings 5 km N of, 38°18'S, 145°11'E, on Avicennia 
marina, 1975, RWR (BRIU973RR); Warrnambool, 38°23'S, 142°20'E, 1886, F R. M. Wilson (NSW 
L4311); Curdies Ck, 38°25'S, 143°00'E, on bush, 1885, F. R. M. Wilson (NSW L4315). South Australia: 
Springton 6-5 km W of, 34°43'S, 139°05'E, on Hakea sp., 1976 JAE (JAE 2261); Mt Barker, 35°06'S, 
138°51'E, on dead twig, 1981, RWR (BRIU2427RR). Tasmania: Trowutta, 4r02'S, 145°05'E, on Pyrrhus 
sp., 1981, GK 391/81; Mt Barrow Chalet, 41°22'S, 147°27'E, on Acacia dealbata, 1969, GB & JC 
(H032547); Upper Esk,41°26'S, 14T44'E, on Acacia dealbata, 1981, GA:60/81;Cressy,41°43'S, 147°02'E, 
on dead Acacia, 1964, JC (H032561); Central Plateau, 4r45'S, 146°50'E, on Athrotaxis cupressoides, 
1980, GK 508/80; Friendly Beach, 42°00'S, 145°15'E, on Acacia sp., 1968, GB & JC (H032332); Shannon 
Lagoon, 42°08'S, 146°51'E, on dead eucalypt, 1980, GK 153/80; Lake Crescent, 42°10'S, 147°09'E, on 
dead Acacia, 1969, GB (H032544); Old Mans Head, 42°12'S, 147°13'E, on dead Acacia, 1964, GB & JC 
(H032333); Ouse, 29 km N of, 42°14'S, 146°44'E, on Acacia sp., 1964, GiB cfe7C(H032557); Dee, 6 km E 
of, 42°17'S, 146°38'E, on Acacia sp., 1970, GB (H032548); Tarraleah, 42°18'S, 146°26'E, on Acacia 
dealbata, 1980, GK 346/80; EUendale, 42°38'S, 146°43'E, on old oak, 1980, GNS (BRIU3231NS); 
Sorrel-Nugent Rd, 42°46'S, 147°39'E, on briar and Acacia sp., 1964, GB (H03:330); Cygnet, Bradleys 
Property, 43°09'S, 147°05'E, on dead willow, 1973, GB (H032322); Port Arthur, 43°13'S, 147°50'E, 1908, 
Mrs Townsend (NSW L4318). 

13. Ramalina whinrayi[^whinrayiP] N. Stevens Plate 9, fig. 7. 

in Lichenologist 18: 187 (1986). Type: Australia, Hogans Island (Bass Strait), 1973, Whinray s.n. (MEL! - 
holotype; divaricatic acid ± nordivaricatic acid). 

Thallus saxicolous, grey-green, caespitose, rigid, up to 1-0 cm high, branching sparse; 
branches arising from the base, branch width 1-0-2-5 mm, subterete to terete, rarely flattened, 
apices attenuate or blunt; surface matt; pseudocyphellae coarse, large, punctiform, numerous; 
holdfast delimited or diffuse when in colonies; soralia absent. 

Apothecia numerous, disc 3-10 mm diam., concave becoming plane to convex, terminal and 
lateral, or at the axils of bifurcating branches, spores ellipsoid, straight 12 x 4-5-5-0 /Ltm. 

Chemistry. Divaricatic acid ± nordivaricatic acid or sekikaic acid 4'-0-demethylsekikaic acid 
and usnic acid. 

Remarks. The anatomy of R. whinrayi differs from that of any of the other Australian taxa, but 
it is similar to the anatomy of/?, capitata (= R. strepsilis) from the northern hemisphere. 

Distribution and habitat. Ramalina whinrayi is restricted in distribution to some of the Bass 
Strait islands and one site on the Tasmanian north coast (Fig. 19). This saxicolous taxon grows 
on granitic boulders at altitudes from 36-84 m above sea-level. The islands where this taxon 
occurs lie along a southeast line - from the Hogan Group, the Kents Group to Craggy Island 
between latitudes 39°12'S and 40°45'S. Ramalina whinrayi also occurs at The Nut, Stanley, 
Tasmania, where it occurs in overhangs on a basaltic cliff facing the ocean and exposed to strong 
winds. 

World distribution. Australia. 

Bass Strait: Hogans Group: Long Islet, 39°12'S, 147°00'E, on granite, 1973, JW (MEL1012965); Hogans I, 
39°13'S, 146°59'E, on granite, 1973, JW (MEL1012948) Type; Kents Group: North East I, 39°27'S, 



G. NELL STEVENS 




Fig. 19 Distribution of R. whinrayi. 

147°25'E, on granite, 1971, JW (MEL1012636); Craggy I, 39°41'S, 147°40'E, on granite, 1972, JW 
(MEL1013031). Tasmania: The Nut, near Stanley, 40°45'S, 145°18'E, on basalt, 1973, GB (H032796). 

14. Ramalina celastri (Sprengel) Krog & Swinscow 

in Norw. J. Bot. 23: 159 (1976). 

Parmelia celastri Sprengel, Syst. Veget. 4 (2): 328 (1827). Type: Cap. B. Spei, Eklon (S! - holotype; FH! - 
isotype; usnic acid). 

subsp. celastri Plate 9, fig. 8; Plate 10, fig. 1; Plate 15, fig. 4. 

Ramalina fraxinea ^yemensis Ach., Lichenogr. Univ.: 602 (1810). Type: Arabia, Forskal, herb. Acharius 

(H! - holotype). - Ramalina yemensis (Ach.) Nyl. in Bull. Soc. Linn. Normandie II, 4: 144 (1870). 
Ramalina ecklonii auct. , non Sprengel. 

Thallus corticolous, rarely saxicolous, pale green to grey-green, erect, subpendulous to 
pendulous, up to 12 cm long, exceptionally to 30 cm; branching initially trichotomous, either 



Plate 10 Thallus growth forms in the Australian Ramalinae. Scale in mm. Fig. 1 R. celastri subsp. celastri 
(common form). Fig. 2 R. celastri subsp. ovalis (broadest form). Fig. 3 R. inflata subsp. perpusilla 
(coastal ecotype). Fig. 4 R. inflata subsp. perpusilla (inland ecotypes). Fig. 5 R. inflata subsp. inflata 
(open-forest ecotype). Fig. 6 R. inflata subsp. inflata (closed-forest ecotype). Fig. 7 R. inflata subsp. 
australis: LHS divaricatic acid specimen RHS sekikaic acid specimen. Fig. 8 R. fissa (branch at centre top 
of photo shows medulla due to splitting apart). 



RAMALINA IN AUSTRALIA 181 

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182 G. NELL STEVENS 

remaining as a three-lobed thallus or thereafter dividing subdichotomously or irregularly, often 
producing densely branched thalli, rarely producing small branchlets at right angles to the 
primary branches; branch width (l-)2-5 mm exceptionally to 20 mm, branches compressed 
ranging from narrow and canaliculate with apices attenuate to broad and flat with apices acute or 
blunt; surface matt, smooth to rugose, membraneous or coarse and ridged with strands of 
chondroid tissue, cracks or holes may appear between these strands; pseudocyphellae usually 
present; holdfast delimited; soralia absent. 

Apothecia numerous, marginal on narrow branches, laminal and marginal on broader 
branches; apothecia occasionally appear on both upper and lower surfaces of the branch; disc 
0-5-2-0 mm diam., plane to convex; margin narrow, entire, smooth or crennate, almost 
disappearing at maturity; spores oval, ellipsoid to slightly fusiform, mainly straight to slightly 
curved, (10-)12-16 x 4-6 /u,m. 

Chemistry. Usnic acid only. \ 

Remarks. The name R. ecklonii (Sprengel) Meyen & Flotow was incorrectly applied to this 
species for a long time. Krog & Swinscow (1976) rejected the name R. ecklonii, applying Article 
69 of the Code, and chose R. sprengelii as a nomen novum for it. Material collected by Ecklon 
from Cape of Good Hope was cited as the holotype. This material is very small, whereas an 
isotype held at FH in Tuckerman herbarium comprises a number of thalli in good condition. All 
of these specimens are narrow and canaliculate in form. Comparison of the type material with 
the narrow ecotype found in Australia showed them to be morphologically similar. 

Specimens bearing this narrow morphology are often confused with R. linearis Swartz, but 
there are distinct differences which easily separate the two (Stevens, 1983ft). Broad thalU which 
belong to R. celastri subsp. celastri resemble the morphology of the holotype of R. yemensis 
(Ach.) Nyl. {R. fraxinea ^yemensis Ach.). This variability in the width of branches caused many 
new taxa to be described, some of which Hue (1890: 265) listed as synonyms ofR. yemensis. 

Weber & Wetmore (1963, 1972) listed the names R. ecklonii Sprengel, R. ecklonii var. 
membranacea (Laurer) Miill. Arg. , R. ecklonii v diX . tenuissima Meyen & Flotow, R. linearis Sw., 
and R. yemensis (Ach.) Nyl. as occurring in AustraHa, but examination of these specimens has 
identified them as belonging to R. celastri subsp. celastri. 

Examination of some South American material indicates several of these species may be 
morphotypes of R. celastri subsp. celastri. R. ecklonii var. lobulifera Malme closely resembles 
the broad morphotype found in Australia. Other species, e.g. R. laevigata and R. continentalis , 
appear to be morphotypes of R. celastri subsp. celastri, but more research into the South 
American taxa is necessary before taxonomic conclusions can be established. The position of R. 
cumanensis Fee in this group of taxa has yet to be determined; narrow canaliculate morphotypes 
in Australia have been found which closely resemble it. Krog & Swinscow (1976) commented 
that a specimen ofR. celastri from Mt Kenya resembled R. cumanensis, and Landron (1972) was 
convinced that R. cumanensis and R. ecklonii were the same species and reduced the latter name 
to synonomy. This problem is still to be resolved. 

Distribution arid habitat. Ramalina celastri subsp. celastri morphotypes extend from the subtro- 
pics (26°S) to the temperate region (38°S) (Fig. 20), with the coastal occurrence more restricted 
in latitude (28°S to 35°S). The inland distribution is extensive in eastern Australia, occurring up 
to 200 km from the coast. It also has a wide altitudinal range, extending from sea-level to the 
mountains. 

Narrow canaliculate morphs occur in montane forest habitats on a variety on tree species. In 
these habitats the rainfall usually exceeds 1500 mm per annum and mists are common 
throughout the year. Broad, flat thalli (some reaching widths of over 10 mm) occur in the drier 
open-forest areas on several phorophytes, species of Acacia being the most common, e.g. A. 
leiocalyx, A. concurrens, A. irrorata, A. orites, A. melanoxylon, and /I. mearnsi. The lichen also 
occurs on Alphitonia excelsior and on the thorny shrubs Citriobatus pauciflora and Capparis 
arborea. 

Ramalina celastri subsp. celastri is usually found growing in association with the fistulose taxa 



RAMALINA IN AUSTRALIA 
TIo" 



183 




Fig. 20 Distribution of R. celastri subsp. celastri • and subsp. ovalis O. 



R. inflata subsp. perpusilla and R. inflata subsp. inflata up to 1000 m, and with R. reducta at 
altitudes above 1300 m. 

World distribution. Pan-subtropical. 

Queensland: Narayen, 25°48'S, 151°10'E, on scrub tree, 1980, N. Gibson (BRIU3287NS); Kingaroy, 3 km 
SW of, 26°34'S, 151°48'E, on dead shrub, 1979, HS (CBG8000062); Jimna, 26°40'S, 152°28'E, on Acacia 
sp., 1980, GNS (BRIU3354NS); Bunya Mts, 26°50'S, 151°40'E, on Acacia sp., 1979, GNS 
(BRIU3289NS); Mt Mee, 27°06'S, 152°42'E, on Citriobatus pauciflora, 1971, RWR (BRIU1919RR); 
Maclagan, 27°10'S, 151°38'E, on Acacia sp., 1979, GNS (BRIU3290NS); Toowoomba, Highfields, 
27°23'S, 151°58'E, on Alphitonia sp., 1983, GNS (BRIU4066NS); Rosewood, 27°39'S, 152°36'E, on 
Alphitonia sp., 1982, GNS (BRIU3559NS); Pittsworth, 20 km W of, 27°45'S, 151°27'E, on Carissa ovata, 
1983, M. Olsen, (BRIU3761NS); Tamborine Mt 27°55'S, 153°12'E, on shrub, 1979, GNS (BRIU3012NS); 
Mt Cordeaux, 28°03'S, 152°23'E, on rainforest tree, 1978, GNS (BRIU3099NS); Warwick, Apex Park, 
28°13'S, 152°02'E, on oak tree, 1980, GNS (BRIU3235NS); Binna Burra, 28°12'S, 153°11'E, on Acacia 
sp., 1979, RWR (BRIU2153RR); O'Reillys, 28°15'S, 153°08'E, on Eugenia sp., 1983, GNS 
(BRIU3735NS); Springbrook, 28°14'S, 153°16'E, on Acacia orrites, 1983, GNS (BRIU3759NS); Wilsons 
Peak,28°15'S, 152°30'E,on^cfldflsp., 1978, GA^5(BRIU3829A.NS); Kelvin Falls, 28°27'S, 152°06'E,on 
Casuarina sp., 1980, NCS (BRIU3286NS). New South Wales: Undercliff Falls, 28°38'S, 152°10'E, on 
Acacia melanoxylon, 1981, GNS (BRIU3652NS); Lismore, 28°48'S, 153°16'E, on Acacia sp., 1976, R. 
Moodie (BRIU2850NS); Yamba, 29°27'S, 153°20'E, on Avicennia marina, 1977, GNS (BRIU2124A.NS); 
Glen Innes, 35 km S of, 30°00'S, 151°40'E, on Acacia sp., 1980, GNS (BRIU3232NS); Clouds Ck, 30°10'S, 
152°35'E, on small tree, 1978, HS (CBG7905564); WoUomombi Falls, 30°32'S, 152°03'E, on Acacia sp., 



184 G. NELL STEVENS 

1982, GNS (BRIU3776NS); Port Macquarie, 31°26'S, 152°54'E, on Avicennia marina, 1977, GNS 
(BRIU2124NS); Walcha Road, 31°25'S, 151°30'E, on Acacia sp., 1982, GNS (BRIU3764NS); Cobak R. 
31°57'S, 151°48'E, on Casuarina sp., 1982, GNS (BRIU3770NS); Gosford, Erina Ck, 33°25'S, 151°20'E, 
on Casuarina glauca, 1978, JAE (JAE 4722); Jenolan Caves, 33°48'S, 150°01'E, on Acacia sp. , 1977, JAE 
(JAE 3180); Wombeyan Caves, 34°19'S, 149°58'E, on dead shrub, 1979, HS (CBG7907795); Nowra, 14 
km N of, 34°48'S, 150°40'E, on tree, 1976, HS (3456); Bundanoon, 34°39'S, 150°18'E, on tree, 1954, M. 
Tindale (MEL9457); Bungendore, 12 km E of, 35°20'S, 149°28'E, on old bones, 1978, JAE (JAE 5465); 
Batemans Bay, 35°45'S, 150°10'E, on Avicennia marina, 1978, GNS (BRIU2170NS); Murunna Pt, 
36°24'S, 150°05'E, on dead coastal shrub, 1978, JAE (JAE 4591); Twofold Bay, 37°05'S, 149°54'E, White 
(MEL 9456). Victoria: Lakes Entrance, 37°53'S, 148°00'E, 1901 (MEL 9465); Frankston, 38°11'S, 
145°03'E, 1900 (MEL 9462); Traralgon, 8 km E of, 38°12'S, 146°32'E, on Melaleuca sp., 1975 RF 
(MEL1013204); Yarragon, Strzelecki Ra. 38°12'S, 146°04'E, on trees, 1973, GB & R. C. Weeks 
(H032292); Trida, Grand Ridge Rd, 38°19'S, 145°58'E, on trees, 1971, GB & R. C. Weeks (H032290); 
Gippsland, Foster 6-4 km NW of, 38°40'S, 146°13'E, on dead wood, 1968, GB (H032297). Tasmania: 
Trowutta, 41°02'S, 145°05'E, on Pyrrhus sp., 1981, G/C 386/81. ^ 

subsp. ovalis (J. D. Hook. & Taylor) N. Stevens, comb. nov. Plate 10, fig. 2; Plate 15, fig. 5. 

Ramalina ovalis J. D. Hook. & Taylor in Lond. J. Bot. 3: 655 (1844). Type: Van Dieman's Land, Gunn s.n. 
(BM! - holotype; usnic acid). - Ramalina calicaris var. ovalis (J. D. Hook. & Taylor) Church. Bab. in J. 
D. Hook., Flora Tasmaniae 3: 344 (1860). - Ramalina ecklonii var. ovalis (J. D. Hook. & Taylor) F. 
Wilson in Pap. Proc. R. Soc. Tasm. 1892: 159 (1893). - Ramalina yemensis var. ovalis (J. D. Hook. & 
Taylor) Zahlbr., Cat. Lich. Univ. 6: 529 (1930). 

Thallus corticolous, green-grey, erect to subpendulous, 2-4(-5) cm long, exceptionally to 8 
cm; branching sparse with one or two branches often constituting the whole thallus; branch 
width 8-20 mm, compressed, flat, apices blunt or rounded, small branchlets may occur at the 
apices; surface matt, membraneous and thin, wrinkled, or thick and ribbed with chondroid 
strands; pseudocyphellae sometimes present; holdfast delimited; soralia absent. 

Apothecia laminal only, numerous, disc to 2-0 mm diam., plane to convex; margin entire, 
thin; spores ellipsoid or slightly fusiform, mostly curved, (10-)12-16 x 4-4-5 jxm. 

Chemistry. Usnic acid only. 

Remarks. There is no record of the exact location of the collection of the type material of R. 
ovalis apart from Van Diemans Land. The obovate form of the branches was commented on by 
Hooker & Taylor (1844) in the type description - 'has the outline of Fucus ovalis' - and this 
morphology is common in Tasmanian material, but mainland specimens are more variable. 

From the taxonomic nomenclature applied to this taxon in the past it appears most authors 
appreciated its affinity to the species R. celastri {= R. yemensis, = R. ecklonii). 

Distribution and habitat. This temperate taxon occupies a broad band across Victoria, South 
AustraUa, and Western Austraha in the latitudes 33°S to 38°S, and is found on the Bass Strait 
islands and in Tasmania along the north and eastern coast, latitudes 40°S-43°S (Fig. 20). It 
virtually replaces R. celastri subsp. celastri in these latitudes and is confined to areas of winter 
rainfall. It grows mainly in open-forest and tall shrubland communities. 

Krog (in lift.) detected the occurrence of this broad taxon in collections ofR. celastri from East 
Africa. Examination of the BM material from Tanzania, Moshi district (1440 m alt.) and 
Uganda (1950-2100 m alt.) showed that thalU with the same morphology as R. celastri subsp. 
ovalis grow in these open-forest areas. This taxon also occurs in New Zealand at comparable 
latitudes to those where it is found in Australia. 

The main phorophytes ofR. celastri subsp. ovalis are Acacia pycnantha, A. melanoxylon, A. 
acuminata, A. longifolia, A. dealbata, and Lycium ferocissimum, Bursaria spinosa, Banksia 
marginata, Casuarina stricta, Callitris sp., and Pyrrhus, Populus, Malus, and Prunus spp. 

World distribution. Africa and Australia. 

New South Wales: Neville, 33°43'S, 149°13'E, on Acacia sp., 1980, GNS (BRIU3302NS); Crookwell, 
34°28'S, 149°29'E, on Acacia sp., 1980, GNS (BRIU3233NS). Victoria: Mario, 37°48'S, 148°32'E, on 
Leucopogon sp., 1980, GNS (BRIU3221NS); Korumburra, Coal Ck, 38°25'S, 145°48'S, on Acacia sp., 



RAMALINA IN AUSTRALIA 185 

1980, GNS (BRIU3248NS); Wilsons Prom, N of, 38°45'S, 146°08'E, on Acacia sp., 1980, GNS 
(BRIU3219NS); Crib Pt, Westernport Bay, 38°22'S, 145°10'E, on Avicennia marina, 1978, GNS 
(BRIU2346NS); Gunnamatta Beach, 38°23'S, 144°45'E, on dune shrub, 1976, RF (MEL1025871); 
Ballarat, 20 km WNW of, 37°30'S, 143°44'E, on Pinus sp. , 1975, E. M. Canning (CBG8100893); Portland, 
38°21'S, 141°36'E, 1947, A. C. Beauglehole (MEL1020517). South Australia: Oodlawirra, 5 km W of, 
32°53'S, 139°03'E, on Muellenbeckia sp., 1969, RWR (BRIU1632RR); Eyre Peninsula, W of Lake 
Newland, 33°22'S, 134°53'E, 1967, N. N. Donner (AD97539274); Yorke Peninsula, Cape Elizabeth, 
34°08'S, 137°27'E, 1969, B. Copley (AD97414422); Port Rickaby, S. of, 34°41'S, 137°29'E, 1974, J. Z. 
Weber (AD97648159); Springton, 14 km SE of, 34°39'S, 139°06'E, on Callitris sp., 1974, JAE (JAE 481); 
Mt Lofty Ra., Clarendon, 35°07'S, 138°38'E, 1968, V. Cruickshank (AD97418421); Trent Hill, 35°36'S, 
138°12'E, on Casuarina sp., 1968, RWR (BRIU1458RR); Naracoorte, 36°58'S, 140°45'E, D. Hunt 
(CBG067824); Kangaroo I, W of Kingscote airport, 35°39'S, 137°38'E, on Eucalyptus sp., R. D. Seppelt 
(MEL1024502). Western Australia: Boyup Brook, 33°50'S, 116°22'E, on Malus sp., 1974, K. M. Allan; 
Lake King, 33°05'S, 119°40'E, on twigs, 1980, D. Richardson (PER000996); Northcliffe, Beedalup Falls 
Rd, 34°38'S, 116°07'E, on dead tree, 1973, B. Muir (N. Sammy Herb); Recherche Archipelago, Peak I, 
34°00'S, 122°50'E, on Acacia acuminata, 1950, /. H. Willis (MEL 10025); Long I, 34°03'S, 12r58'E, on 
Melaleuca sp., 1950, /. H. Willis (MEL 10020). Bass Strait: Curtis I, 39°29'S, 146°39'E, 1971, McCoy 
Expedition, (MEL40274); Furneaux Group: Forsyth I, 40°38'S, 148°25'E, on Acacia sp., 1976, JW 
(MEL1516801). Tasmania: Stanley, The Nut, 40°45'S, 145°15'E, on Bursariaspinosa, 1965,7. H. Willis 
(MEL 7588); Devonport airport, 41°10'S, 145°20'E, on Poplar sp., 1980, GNS (BRIU3236); Avoca, 
41°48'S, 147°40'E, on Prunus sp., 1978, GK 2/78; Sorrell-Nugent Rd, 42°46'S, 147°39'E, on briars, 1964, 
GB&7C(H032314). 

15. Ramalina inflata (J. D. Hook. & Taylor) J. D. Hook. & Taylor 

in J. D. Hook., Flora Antarctica 1: 194 (1845). 

Cetraria inflata 3. D. Hook. & Taylor in Lond. J. Bot. 3: 646 (1844). Type: Lord Auckland Islands, Hooker 

s.n. (FH! - holotype; BM! -isotype; divaricatic acid and usnic acid) .- Fistulariella inflata (J. D. Hook. & 

Taylor) Bowler & Rundel in Mycotaxon 6: 195 (1977). 

subsp. inttata Plate 10, figs 5-6; Plate 14, fig. 5. 

Ramalina tasmanica Nyl. in Bull. Soc. Linn. Normandie II, 4: 162 (1870) type: Australia, Tasmania, /. D. 

Hooker s.n. (H! - holotype [H-NYL 36849 ex hb. Churchill Babington] ; divaricatic acid and usnic acid). 

Plate 14, fig. 6. 
Ramalina knightiana Zahlbr., Cat. Lich. Univ. 6: 494 (1930); Ramalina subgeniculata Knight in Trans. 

Linn. Soc. Lond. (Bot.) 2: 50 (1882), non Nyl. (1870) (Art 64.1). Type: Australia, New South Wales, 

near Sydney, Knight s.n. (WELT! - holotype; divaricatic acid, sekikaic acid, and usnic acid). 

Thallus corticolous, green to pale green, caespitose, erect l-2(-3) cm high; branching 
subdichotomous or irregular; branch width 1-3 mm, branches hollow, terete, inflated and 
perforate; perforations round to elongate, medullary hyphae continuous, loosely woven across 
central cavity or compressed against the inner cortex wall either loosely or densely; surface matt 
to shiny, smooth, rarely pseudocyphellate; holdfast delimited; soralia absent. 

Apothecia common, terminal on main branches and on short subapical lateral branches, often 
spurred; disc 2-5 mm diam. , always concave initially, innate at branch apices, becoming plane at 
maturity; margin entire, thin often indistinct; spores broadly ellipsoid, straight or curved, 12-16 
X 4-5 (-6) /Am. 

Chemistry. Divaricatic acid, ± nordivaricatic acid, ± sekikaic acid, and usnic acid. 

Remarks. Sterile specimens of this taxon are usually larger than fertile thalli, occurring either as 
greatly inflated, pulvinate, thalli or having elongate branches with perforations evenly distrib- 
uted along the lower surface and extending to the attenuate apices. 

A small morphotype of R. inflata subsp. inflata found in Tasmania was named R. tasmanica 
Nyl. ; the type material contains divaricatic acid. Subsequent collections from Tasmania bearing 
this name have been found to contain salazinic acid and belong to R. fissa. The type material of 
R. knightiana Zahlbr. is a very finely branched specimen of R. inflata subsp. inflata collected in 
the neighbourhood of Sydney; its morphology closely resembles the New Zealand species R. 
geniculata and was originally named R. subgeniculata by Knight. 



186 G. NELL STEVENS 

The synonomy of R. inflata and R. geniculata has been the subject of debate by several 
Hchenologists (Howe, 1913-14; Landron, 1972; Bowler & Rundel, 1977). An examination of 
the types of both of these species during the present study has shown them to contain the same 
acid (divaricatic) but to differ morphologically. More material from New Zealand (from which 
country the type of R. geniculata was collected) would need to be examined to evaluate the 
amount of phenotypic plasticity displayed by this taxon and to compare the morphotypes with R. 
inflata ecophenotypes, in order to establish if both taxa intergrade with one another. 

In many respects the morphology and habitat requirements of the East African taxon R. 
calcarata Krog & Swinscow come close to the Australian specimens of R. inflata subsp. inflata, 
but the presence of salazinic acid in the hymenium of the former taxon distinguishes the two. A 
tan coloration on the lower surface of some specimens of R. calcarata was mentioned by Krog & 
Swinscow (1974) and this colour occasionally occurs on the Australian material of R. inflata 
subsp. inflata. However discoloration of the thallus is not common and is not regarded as of any 
taxonomic significance. The original protologue oiR. inflata contains the comment: 'sometimes 
stained with dark blood red', but this colour is rarely evident on specimens. However black 
patches on the thallus of R. pusilla Le Prev. ex Duby., have been commented upon by Krog & 
0sthagen(198O). 

Distribution and habitat. The distribution of R. inflata subsp. inflata (Figs 21 , 22) extends along 
the warm to cool temperate section of the eastern and south-eastern Australian coastal belt and 
into the cool/cold temperate region of Tasmania. The most northern occurrence of this taxon is 





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RAMALINA IN AUSTRALIA 



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in the mountains at elevations of 1000 m in the area 28°-30°S latitude. This is an area where 
intermediates between the morphology of the rain-forest ecotype of R. inflata subsp. perpusilla 
and the morphology of R. inflata subsp. inflata occur. At 30°S latitude (WoUomombi Falls, 
elevation 1000 m) R. inflata subsp. inflata grows on Acacia in open forest communities growing 
on metamorphic rock, and the thalli are typical; at 32°S latitude (Barrington Tops, a basalt 
range) thalli collected from rain-forest areas were difficult to place in either taxon, indicating 
that the microhabitat as well as climatic conditions may be influencing these two subspecies. 
The most common phorophytes for R. inflata subsp. inflata are Acacia and Casuarina spp. 

World distribution. South Africa, Australia, Auckland Island, Brazil, and Venezuela. [Uncon- 
firmed reports are from China (Zahlbruckner, 1930), North America (Howe, 1914), West Indies 
(Landron, 1972), and Hawaii (Magnusson, 1955)]. 

New South Wales: Undercliff Falls, 28°38'S, 152°10'E, on Acacia sp., 1980, GNS (BRIU3028A.NS); 
WoUomombi Falls, 30°30'S, 152°03'E, on Acacia sp., 1982, GNS (BRIU3639NS); Barrington Tops, 
32°00'S, 151°30'E, on rain-forest tree, 1982, GNS (BRIU3636NS); Scone, 12 km W of, 32°05'S, 150°55'E, 
on bush, 1982, GNS (BRIU3635NS); Tuncurry, 32°10'S, 152°27'E, on Casuarina sp., 1982, GNS 
(BRIU3648NS); Erina Ck, 33°25'S, 151°21'E, on Casuarina glauca, 1978, JAE (JAE4705); Towra Pt, 
34°02'S, 151°13'E, on Casuarina sp., 1977, CS (BRIU63CS); Barrengarry Mt, 34°41'S, 150°34'E, on tree, 
1976, JAE (JAE2714); Currowan State Forest, 35°39'S, 150°08'E, on Acacia sp., 1977, JAE (JAE3576); 
Batemans Bay, 35°42'S, 150°10'E, 1970, ED (0); Batemans Bay, on Avicennia marina, 1978, GNS 
(BRIU2172NS);Wallaga Lake, 36°23'S,150°03'E, on Co^uannflsp.n., 1978, /^E (JAE 4543); Spring Ck 



188 G. NELL STEVENS 

Gap, 37°30'S, 148°20'E, on Acacia sp., 1978, JAE (JAE5156); Mario, 37°48'S, 148°32'E, on Acacia sp., 
1978, GNS (BRIU3282NS); Cape Conran, 37°49'S, 148°45'E, on Leptospermum sp., 1983, GNS 
(BRIU3676NS); Rodondo I, 39°15'S, 146°23'E, 1970, J. Kirkpatrick (MEL39969); Cape Patterson, 
38°40'S, 145°35'E, on Acacia pycnantha, 1963, RF & Smith 3413; French I, Tankerton Rd, 38°20'S, 
145°20'E, S. Filson (MEL1026170); HealesviUe, 37°39'S, 145°32'E, on twigs, 1974, RB (MEL1013089); Mt 
Richmond Nat. Park, 38°18'S, 141°30'E, 1978, AA (BRIU4312NS). Bass Strait: Kents Group: Erith I, 
39°28'S, 147°13'E, on Beyeria leschenaultia, 1970, JW (MEL1012899). Tasmania: Rocky Cape, 40°51'S, 
145°31'E, 1975, D & M. Cook (H032566); Yolla, 4r09'S, 145°40'E, on Malm sp., 1980, GNS 
(BRIU3239NS); Mt Barrow, 41°23'S, 147°25'E, on Acadasp., 1966, GB (H032678); Liffey Falls, 41°41'S, 
146°47'E, on Acacia sp., 1969, GB & JC (H032752); Tarraleah, 42°15'S, 146°26'E, on Acacia dealbata, 
1980, GK 347/80; Mt Faulkner, 42°48'S, 147°12'E, on Acacia sp. , 1965, GB & JC (H032762). 

subsp. perpusilla (Stirton) N. Stevens, comb. nov. Plate 10, figs 3-4; Plate 14, fig. 3. 

Ramalina perpusilla Stirton in Proc. R. Soc. Vict. 17: 68 (1881). Type: Australia, Queensland, Brisbane, 

Bailey 113 (BRI! - lectotype; herein designated, left hand specimen; divaricatic acid and usnic acid). 
Ramalina minuscula var. alba C. Knight ex Shirley in Proc. R. Soc. Qd 5: 103 (1888). Type: Australia, 

Queensland, Mt Perry, Keys 91 (WELT! - lectotype; sekikaic acid and usnic acid). Plate 15, fig. 7. 
Ramalina geniculata var. compacta Miill. Arg. in Bull. Herb. Boissier 4: 88 (1896). Type: Australia, 

Queensland, Toowoomba, Hartmann s.n. (G! - lectotype designated here; divaricatic acid and usnic 

acid). 

Thallus corticolous, green to pale green, often with white lower surface, caespitose, erect, 
l-2(-3) cm high; branching subdichotomous, branch width 0-5-2-0 mm, varying in the amount 
of inflation, terete to almost flat, sparingly to moderately perforate, the perforations on the 
lower surface only, small and round, extending to the apices on sterile branches; medullary 
hyphae sparse and adhering to the inner cortex wall either loosely or densely compacted in 
totally inflated specimens or thick and felty in compressed thalli; surface matt or shiny, smooth 
to rugose; very rarely pseudocyphellate; holdfast delimited or diffuse; soralia absent. 

Apothecia sparse to numerous, subterminal on geniculate branches or terminal on main and 
short lateral branches, spurred, or in the axils of bifurcating branches, disc usually less than 2-0 
mm diam., plane to convex; margin entire, well defined, rarely indistinct; spores ellipsoid, 
straight, rarely curved, (8-)10-14(-16) x 4-5 /xm. 

Chemistry. Divaricatic acid ± trace sekikaic acid, ± nordivaricatic acid or sekikaic acid only, or 
rarely sekikaic acid, 4'-0-demethylsekikaic acid, an unknown and usnic acid. 

Remarks. The type material of R. inflata subsp. perpusilla comprises four specimens with the 
number 113 apparently covering three of the specimens and the number 259 covering the fourth 
specimen. The left hand specimen contains divaricatic acid and is the most likely single specimen 
that number 113 could refer to (Plate 14, fig. 3). The middle and right hand top specimens 
contain sekikaic acid as does specimen 259. The left-hand specimen has been selected as 
lectotype for this taxon because it appears to conform best with the original protologue. As the 
majority of specimens collected during this study contain divaricatic acid as the major acid, the 
choice of a lectotype which contained divaricatic acid seemed appropriate. 

The only locality given for the type collection is Brisbane River. In describing this species 
later, Shirley (1888) gave the information 'Hocking's Nursery, Brisbane River'. It is not clear 
however, whether the specimens were growing on native trees in the area, or on the mangroves, 
or were found on one of the nursery trees, which could have come from elsewhere. Specimens 
which produce sekikaic acid only have not been found on mangroves; they are usually found in 
dry inland situations, which woyld indicate that the type material possibly grew on a plant which 
had come from another location. This problem remains unsolved and the true type locality is not 
known. 

Ramalina minuscula var. alba was collected at Mt Perry, a dry open-forest area in southern 
Queensland. The type material consists of 11 carded specimens all of which contain sekikaic 
acid. 

Ramalina geniculata var. compacta was collected in the Toowoomba area. The material 
comprises six specimens on a card (a, b, c, d, e, f); b and d contain divaricatic acid whereas a, c, e 



RAMALINA IN AUSTRALIA 189 

and f contain sekikaic acid. Specimen b is designated lectotype as it conforms best with the 
original protologue and contains divaricatic acid, consistent with the type of R. inflata subsp. 
perpusilla with which it is synonymous. As Toowoomba is an inland region about 130 km from 
the coast, the presence of sekikaic acid specimens as well as divaricatic acid specimens is not 
unusual because fresh collections made during the present study from inland areas always 
contained sekikaic acid thalli as well as divaricatic acid thalli. The sekikaic acid race has not been 
given any taxonomic rank because of the co-existence of both divaricatic and sekikaic acids in 
manyof thethaUi. 

The morphology of the East African taxon R. consanguinea Miill Arg. from the mangroves 
resembles R, inflata subsp. perpusilla specimens from the mangrove communities. Both produce 
apothecia which are plane to convex but never concave, and the former taxon has short slender 
side branches and spinules on the main branches, a feature sometimes present on Australian 
specimens. Ramalina consanguinea, however, contains both divaricatic acid and norstictic acid 
and produces large spores 20-25(-28) x 4-5(-6) ^tm, which sets it apart from R. inflata subsp. 
perpusilla. 

Ramalina pumila Mont., an Asian taxon, appears to be close to the Australian taxon 
morphologically, but its chemistry (evernic acid with trace amounts of sekikaic acid) sets the two 
taxa apart. 

Distribution and habitat. Ramalina inflata subsp. perpusilla has a distribution range from the 
tropics to the subtropics (Figs 21, 22). The highly variable morphology found in this taxon over 
this range is ecophenotypic. Three distinct habitats are considered to explain the ecotypes that 
occur, e.g. (1) the mangrove habitat, (2) the open-forest habitat, and (3) the rain-forest 
(closed-forest) habitat. 

(1) The mangrove habitat: Specimens of R. inflata subsp. perpusilla collected from mangrove 
communities along the Queensland coastline from Hallidays Bay (latitude 20°56'S) to Moreton 
Bay (latitude 28°S) are usually small and caespitose with elongate perforations spread irregu- 
larly along the lower surface; short spinules are sometimes present on the main branches and the 
margins of the apothecia are very distinct. North of latitude 23°50'S (Gladstone), 80% of the 
specimens were found to contain only divaricatic acid whereas 45% of specimens south of 
Gladstone contained divaricatic acid and trace amounts of either sekikaic acid or nordivaricatic 
acid. 

The main mangroves which act as phorophytes for this taxon are Ceriops tagal and Rhiz- 
ophora stylosa, and when these mangroves reach their southern distribution limit at latitude 
28°S, R. inflata subsp. perpusilla ceases to occur along the coastline. 

(2) The open-forest habitat (including disturbed habitats) : These dry inland areas lie between 
the maritime and the montane-rain-forest environments and collections of R. inflata subsp. 
perpusilla from such habitats are scattered from latitudes 18°S-27°S. The thalli are very small 
with a maximum size of less than 1 -0 cm high ; they are usually green or pale green with a smooth , 
shiny surface, wholly inflated; perforations are round and evenly distributed on the lower 
surface of the thallus; the branches are either tightly packed together with numerous terminal 
apothecia covering the thallus (Plate 10, fig. 3) or are palmate at the base, dividing into several 
branches which bear subterminal apothecia or apothecia situated in the axils of bifurcating 
branches. 

In this open-forest habitat more than 50% of the population contains sekikaic acid only with 
the remainder containing divaricatic acid ± trace amounts of sekikaic acid. The most common 
phorophytes in such habitats are the small thorny shrubs Citriobatus pauciflora and Acalypha 
capillipes. They grow on basaltic soils in open areas experiencing full sunlight and varying 
humidity; areas which in some respects resemble the open mangrove community where Ceriops 
tagal grows. 

(3) The rain-forest habitat: Pockets of rain-forest where this particular ecotype occurs extend 
from the tropics to the subtropics along the Eastern Highlands (viz. Atherton Tableland (17°S), 
Mt Surprise (19°S), Bunya Mts (27°S), McPherson Ra. (28°S), and Dorrigo Plateau (30°S). 
Specimens of R. inflata subsp. perpusilla from this habitat tend to be loosely branched, 2-3 cm 



190 G. NELL STEVENS 

long, with a pale lower surface; branching is subdichotomous with the branches less inflated, 
even partially compressed, and more rugose than those of the taxa from more exposed habitats; 
the perforations are small, slightly elongate and sparse; the apothecia are usually terminal and 
subterminal, with some tendency to be innate when immature but becoming plane at maturity. 

The majority of the rain-forest specimens contain only divaricatic acid, with a small pro- 
portion producing trace amounts of either sekikaic acid or nordivaricatic acid in addition to 
divaricatic acid. The main phorophytes are various species of rain-forest tree, including 
Araucaria cunninghamii and Eugenia sp., and also Acacia melanoxylon and A. orrites which 
fringe the rain-forest borders. 

Although this taxon occupies three distinct habitats, there is total intergradation of morpho- 
types from one environment to another which prevented any division of the group being made 
based on microhabitat morphology. Chemistry was not able to be used taxonomically, as 
variation from divaricatic acid only in the thallus to divaricatic acid + trace sekikaic acid to 
sekikaic acid only was found to occur. 

One inflated specimen collected from Lake Eacham, Atherton Tableland (JAE 2427), has not 
been included in this taxon; it is a large specimen (4 cm long), partially inflated, partially 
compressed (which is normal for the rain-forest habitat in which it grew) with white punctiform 
pseudocyphellae along the branches. The acids present are sekikaic, homosekikaic, 4'-0- 
demethylsekikaic, and an unknown. This chemistry differs from that produced by the majority 
of the taxa in this group but it is similar to the acids found in a few inland specimens of R. inflata 
subsp. perpusilla. The presence of white pseudocyphellae, however, is quite distinct. As no 
other specimens have been collected from Atherton Tableland, or elsewhere, the specimen 
remains unnamed at present. 

World distribution. Australia. 

Queensland: Tinaroo Dam, 17°16'S, 145°34'E, on Araucaria cunninghamii, 1983, GNS (BRIU4167NS); 
Forty Mile Scrub, 18°03'S, 144°53'E, on thorny shrub, 1983, GA^5(BRIU4114NS);BurdekinR, Big Bend, 
19°15'S, 146°12'E, on thorny bush, 1983, GNS (BRIU4154NS); Hallidays Bay, 20°56'S, 149°01'E, on 
Ceriops tagal, 1983, GNS (BRIU4118); Eimeo Beach, 21°05'S, 149°13'E, on Ceriops tagal, 1976, GNS 
(BRIU2221NS); St Lawrence, 22°20'S, 149°32'E, on Ceriops tagal, 1983, GNS (BRIU4105NS); Marl- 
borough, 22°50'S, 149°51'E, on thorny bush, 1983, M. Olsen (BRIU4157NS); Port Alma, 23°35'S, 
150°42'E, on Rhizophora stylosa, 1975, RWR (BRIU696RR); Gladstone, 23°50'S, 151°16'E, on Ceriops 
tagal, 1980, GNS (BRIU3653NS); Cania Gorge, 24°38'S, 150°56'E, on Casuarina sp., 1984, NCS 
(BRIU4343NS); Bundaberg, The Hummock, 24°51'S, 152°26'E, rainforest tree, 1977, /. R. Telford 
(CBG7702460); Goodnight Scrub, 25°15'S, 151°50'E, on thorny bush, 1981, P. Forster (BRIU3640NS); 
Toogoom, 25°15'S, 152°40'E, on Callitris sp., 1983, GNS (BRIU4150NS); Coalston Lakes, 25°35'S, 
151°55'E, on thorny bush, 1980, RWR (BRIU3586RR); Narayan, 25°48'S, 151°10'E, on shrub, 1980, RWR 
(BRIU2197RR); Barambah Ck, 25°40'S, 15r45'E, on shrub, 1982, NCS (BRIU3606NS); Mt Blandy, 
25°40'S, 151°38'E, on thorny bush, 1980, P. Forster (BRIU3641NS); Lake Cootharaba, 26°18'S, 152°58'E, 
on Bruguiera sp., 1978, GNS (BRIU3620NS); Maleny, 26°46'S, 152°51'E, on Acacia sp., 1982, GNS 
(BRIU3568NS); Bunya Mts, 26°50'S, 151°40'E, on rainforest shrub, 1979, GNS (BRIU3009NS); Mt Mee, 
27°06'S, 152°42'E, on Citriobatuspauciflora, 1979, GNS (BRIU3654NS); Stradbroke 1, 27°25'S, 153°27'E, 
on Lumnitzera racemosa, 1977, GNS (BRIU2118NS); Lake Broadwater Park, 27°21'S, 151°05'E, 1983, B. 
Ballingall 981; Worlds End Pocket, 27°37'S, 152°55'E, on Acalypha capillipes, 1983, P. Forster 
(BRIU3709NS); Toowoomba, Cooby Ck Rd, 27°23'S, 151°38'E, on Citriobatus pauciflora, 1983, GNS 
(BRIU3685NS); Pittsworth, 27°45'S, 151°38'E, on Carissa ovata, 1983, M. Olsen (BRIU3762NS); 
Coochiemudlo I, 27°35'S, 153°20'E, on Rhizophora stylosa, 1981, GNS (BRIU3623NS); Coomera I, 
27°55'S, 153°25'E, on Rhizophora stylosa, 1979, GNS (BRIU3649NS); Tamborine Mt, 27°55'S, 153°12'E, 
on rain-forest tree, 1979, GiV5 (BRIU3010NS);MtCordeaux,28°03'S, 152°23'E, on rain-forest tree, 1979, 
GNS (BRIU3029NS); Springbrook, 28°15'S, 153°16'E, on rain-forest tree, 1982, RWR (BRIU2567RR); 
O'Reillys, 28°15'S, 153°08'E, on rain-forest tree, 1983, 'GA^5 (BRIU3718NS); The Head, 28°18'S, 
152°27'E, on rain-forest tree, 1980, GNS (BRIU3621NS); Kelvin Falls, 28°23'S, 152°06'E, on Casuarina 
sp., 1980, NCS (BRIU3286A.NS). New South Wales: Wiangaree, 28°25'S, 153°08'E, on rainforest tree, 
1978, GNS (BRIU3028NS); Undercliff Falls, 28°38'S, 152°10'E, on Acacia sp., 1980, GNS 
(BRIU3651NS); Cherry Tree State Forest, 28°54'S, 153°01'E, on vine, 1983, B. Ballingall 
(BRIU3646NS); Clouds Ck, State Forest, 30°10'S, 152°35'E, on shrub, 1978, HS (CBG9705565); Mt 
Dorrigo, 30°20'S, 152°40'E, on rain-forest tree, JAE (JAE 3447). 



RAMALINA IN AUSTRALIA 191 

subsp. australisN. Stevens, subsp. nov. Plate 10, fig. 7. 

Thallus corticolus et saxicolus, virido-griseus parvus caespitosus rigidus erectus, usque ad 1-0 cm altus, 
ramificatione nulla vel sparsa, ramis e basi exorientibus; rami lati usque ad 10 mm, valde cavi; superficies 
impolita, laevigata et tenui vel rugosa et crassi; soralia nulla. Apothecia numerosa, terminalia, non 
calcarata, primum immersa, demum plana; discus usque ad 10 mm in diametro; sporae late ellipsoideae, 
curvatae aut rectae, (10-)13-14 x 4-5(-6) /xm. Acidum divaricaticum ± sekikaicum aut sekikaicum et 
usnicum continens. 

Holotype: Australia, Western Australia, Moora-Perth Road on Acacia sp. 6 August 1984, G. N. Stevens 
4315 (MEL 1048089!). 

Thallus corticolous and saxicolous, greenish-grey, small caespitose, rigid, erect, usually to 1-0 
cm high, branching sparse, at the base only; branches broad, up to 10 mm wide, highly inflated, 
perforations large, round to elongate and often splitting apart revealing a sparse medulla and 
internal chondroid strands forming a reticulate fibrose surface on the inner cortex wall; surface 
matt, smooth and thin to rugose and extremely coarse, rarely pseudocyphellate; holdfast 
delimited; soralia absent, although some rock specimens with eroded apices appeared pseudo- 
sorediate. 

Apothecia numerous, terminal, not spurred, sometimes lamina! ; disc up to 5-0 mm, excep- 
tionally to 10 mm diam., concave initially, remaining so or becoming plane; margin entire, 
inrolled; spores broadly ellipsoid, mainly curved, some straight, (10-)12-14 x 4-5(-6) /am. 

Chemistry. Divaricatic acid ± sekikaic acid or sekikaic acid only and usnic acid. 

Remarks. Two acid races occur in R. inflata subsp. australis, (1) a divaricatic acid race and (2) a 
sekikaic acid race. The only name which has been applied to this taxon in the past is R. pusilla, a 
northern hemisphere Ramalina which contains sekikaic acid with salazinic acid in small amounts 
(Krog & Swinscow, 1974). 

Krog & 0sthagen (1980) reported that one specimen of R. pusilla from Hierro, Canary 
Islands, contained salazinic acid only, which indicates that some relationship could exist 
between the taxa which produce sekikaic acid and R. fissa, which produces salazinic acid only. 
Partly because of this, R. fissa is placed in the R. inflata complex. Morphologically the sekikaic 
acid thalli of R. inflata subsp. australis seem closer to the salazinic acid R. fissa thalli than to the 
divaricatic acid thalli. However, the presence of divaricatic acid and sekikaic acid in the same 
thallus shows the close connection between the two depside races, thus preventing the sekikaic 
acid race being separated into a separate taxon from the divaricatic acid race. 

See Table 11 for a comparison of the three subspecies. 

Distribution and habitat. Ramalina inflata subsp. australis occurs in the temperate region of 
mainland Australia and in the drier north and east coastal areas of cool-temperate Tasmania 
(Figs 21, 22). The two chemical races which occur in this taxon are sympatric over most of their 
range, although only the divaricatic acid race occurs coastally . No sekikaic acid thalli were found 
in Tasmania. 

Within its range R. inflata subsp. australis exhibits a variety of ecophenotypes. In the drier 
inland regions of South AustraUa extremely small thalli are found bearing large terminal 
apothecia which obscure the branches (Plate 10, fig. 7). Both divaricatic acid and sekikaic acid 
races occur in such areas, with a predominance of the latter race. In some habitats this taxon 
produces a single inflated branch resembling a small round balloon. 

The divaricatic acid race also occurs on rock substrates, e.g. sandstone (Mt Arapiles and the 
Grampians) and on trachyte (Hanging Rock). Although the morphology of the saxicolous thaUi 
differs from the corticolous thalli, the differences are regarded as the result of change in 
substrate rather than genotypic, e.g. the thalli are much reduced in size, occurring as smaU 
button-like growth forms in colonies; they produce prominent apothecia between sterile 
branches. This rock morphotype resembles the northern hemisphere saxicolous taxon R. 
mediterranea Magnusson, but the Mediterranean species contains evernic acid. It could be 
argued that this change in substrate and morphology warrants the erection of a new species for 
this rock morphotype. However, it is regarded as an ecophenotype oiR. inflata subsp. australis. 



192 



G. NELL STEVENS 






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RAMALINA IN AUSTRALIA 193 

Ramalina inflata subsp. australis is mainly corticolous and grows on various tree species. 
Exotic phorophytes are Pinus, Populus, and Malus; native phorophytes include Acacia, 
Casuarina, Leucopogon, Sind Melaleuca spp. 

World distribution: Australia. 

New South Wales: Batemans Bay, 35°42'S, 150°10'E, on Avicennia marina, 1978, GNS (BRIU2172NS); 
Murunna Pt, 36°23'S, 150°03'E, on coastal shrub, 1978, JAE (JAE 4592). Victoria: Mario, 37°48'S, 
148°32'E, on Melaleuca sp., 1978, GNS (BRIU3201NS); Wilsons Prom., N of, 38°45'S, 146°08'E, on 
Acacia sp., 1980, GNS (BRIU3218NS); Mt Macedon, Hanging Rock, 37°23'S, 144°36'E, on trachyte, 
1976, GNS (BRIU3128NS); Spargo Ck, Korwerigaboora Springs, 37°30'S, 144°00'E, 1965, M. Allender 
(MEL1026135); Triodea Hill, 37°57'S, 141°19'E, on Banksia sp., 1965, /?F (MEL1026193); Drik Drik, 
38°00'S, 141°18'E, 1952,^. C. Beauglehole (MEL544971); Rocket Lake, 34°39'S, 141°48'E, 1970, ED (0); 
Big Desert, 35°22'S, 141°13'E, 1970, ED (0); The Grampians, 37°08'S, 142°26'E, on rock, RF 
(MEL1026047); Dimboola Nat. Park, 36°20'S, 142°01'E, 1949, A. C. Beauglehole (MEL1020493). South 
Australia: Oodlawirra, 5 km W of, 32°53'S, 139°04'E, on Muellenbeckia sp., 1969, RWR (AD97649267); 
Yunta, 40 km S of, 32°59'S, 138°50'E, 1971, N. N. Donner (AD97407411); Blanchetown, 13 km W of, 
34°21'S, 139°29'E, 1973,/. Z. We6er(AD97647391);Naracoorte,25kmSEof,36°58'S, 140°45'E, 1969,/. 
Z. Weber (AD97417116); York Peninsula, Treasure Cove, 35°00'S, 137°25'E, on dead shrub, 1979, JAE 
(JAE 6367); Eyre Peninsula, Streaky Bay, 40 km SE of, 32°48'S, 134°13'E, 1967, N. N. Donner 
(AD97648422); Colona Homestead, 31°38'S, 132°04'E, 1947, /. H. Willis (MEL1026176); Yalata Road 
House, 5 km W of, 3r39'S, 129°03'E, on bushes, 1980, AA (BRIU4345NS). Western Australia: Three 
Springs, 5 km from, 29°32'S, 115°46'E, 1969, on edge of salt lake, R. A. Saffrey (PER000427); Winchester, 
8 km W of, 29°46'S, 115°56'E, on tree, 1966, P. G. Wilson (PER000426); Toodyay-Bindoon Rd, 31°25'S, 
116°15'E, on Acacia sp., 1984, GNS (BRIU4316NS); Pt Mt Henry, Perth, 32°02'S, 116°05'E, on Dryanda 
sessilis, 1970, N. Sammy; Beverley, 17 km S of, 32°10'S, 116°56'E, on Acacia sp., 1984, GNS 
(BRIU4319NS); Hyden, The Humps, 32°19'S, 118°57'E, on granite rock, 1972, N. Sammy; Kondinin 
Forestry Reserve, 32°30'S, 118°24'E, on bushes, 1967, GB dll^dd; Cranbrook 17 km N of, 34°15'S, 
117°37'E, on Casuarina sp. , 1984, GNS (BRIU4314NS); Lake Muir, E side, 34°29'S, 116°40'E, 1980, E. A. 
Griffin (PER001630); Albany-Borden off Chillinup Rd, 34°20'S, 118°12'E, on Melaleuca sp., 1970, N. 
Sammy; Lake Biddy, near Lake Grace, 33°00'S, 118°56'E, on tree, 1976, A. C. Smith (N. Sammy); 
Gairdner R., 34°14'S, 119°17'E, on Hakea sp., 1980, D. Richardson (PER000941); Cape Arid Nat. Park, 
34°02'S, 123°09'E, on Hakea clovata, 1971, R. D. Royce (PER000428). Tasmania: Rocky Cape, 40°53'S, 
145°30'E, on quartzite rock, 1980, GK 743/80; Ansons Bay Nth, 41°02'S, 148°16'E, on Banksia sp., 1975, 
GB (H032680); Moores Hill, Beaconsfield, 4ri3'S, 146°45'E, on Acacia dealbata, 1980, GK 216/80; 
Buckland, Pitts Hill, 42°36'S, 147°43'E, on Bursaris spinosa, 1980, G/C 450/80; Hobart, Gunners Quoin, 
42°54'S, 147°19'E, on dolerite rock, 1980, GX 600/80. 

16. Ramalina fissa (Miill. Arg.) Vainio Plate 10, fig. 8. 

in Mem. Herb. BoissierS: 2 (1900). 

Ramalina inflata var. fissa Miill. Arg. in Flora, Jena 71: 203 (1888). Type: Australia, King Island, Bass 

Strait, 1888, French s.n. (G! - holotype; salazinic acid and usnic acid). Plate 14, fig. 4. 
Ramalina brevis var. brevissima F. Wilson in Pap. Proc. R. Soc. Tasm. 1892: 159 (1893). Type: Austraha, 

Tasmania, Ulverstone, Antill Ponds, Weymouth s.n. (not located). - Ramalina fraxinea var. brevissima 

(F. Wilson) Zahlbr., Cat. Lich. Univ. 6: 486 (1930). 

Thallus corticolous and saxicolous, greenish-grey, erect, rigid to flaccid, inflated and shrubby 
or flattened and almost foliose, 1-3 cm high; branching from the base only; branches variable in 
width, often bursting open to reveal the medulla, giving a false effect of gross inflation, with 
flattening of the thallus the areas directly below the apothecia remain the only part of the thallus 
with a central cavity; perforations when present, large, round to elongate, splitting to reveal the 
medulla; medulla loosely woven, sparse, adhering in patches to the inner cortex wall, the wall 
often marked with reticulate strands of chondroid tissue; sometimes pseudocyphellate; holdfast 
delimited; soraUa absent. 

Apothecia numerous, terminal, not spurred; sometimes laminal; disc 2-5(-8) mm diam., 
initially concave becoming flat; margin entire, inroUed; spores ellipsoid, mainly curved, some 
straight, (10-)12-16 X 4-5(-6) pm. 

Chemistry. Salazinic acid and usnic acid. 



194 G. NELL STEVENS 

Remarks. Ramalina brevis var. brevissima F. Wilson has been placed in the synonymy. 
Although the type material was not located (it is possibly lost), other specimens identified by 
Wilson as R. brevis var. brevissima contain salazinic acid and are small caespitose morphotypes 
oiR.fissa. 

Wilson identified large coarse morphotypes of this taxon as R. fraxinea f. platyna Nyl. 
Material held at G under the latter name consists of two thalh which contain divaricatic acid and 
one which contains salazinic acid. Magnusson examined these specimens and found different 
colour reactions, K— (divaricatic acid), and K + y (salazinic acid). These specimens have been 
found to belong to R. inflata subsp. australis and R. fissa. 

Ramalina subpusilla comes near to R. fissa because of its similar chemistry, but morphologi- 
cally it differs by producing spurred apothecia, a characteristic not found in the latter taxon. 
Ramalina calcarata Krog & Swinscow contains divaricatic acid and small amounts of salazinic 
acid in the hymenium; Krog & Swinscow (1974) compared it to R. fissa because of the 
reticulate-fibrose underside of both and the straight to reniform spores produced by both, but 
the absence of spurred apothecia in the latter taxon separates the two. Galloway (1985) recorded 
that R. geniculata (a New Zealand species often confused with R. infiata) contains sekikaic and 
salazinic (tr.) acids; but the type material (held at FH) contains divaricatic acid. Such variation in 
acid content of a single species adds weight to the suggestion that in Australia there is an 
evolutionary relationship between the depside taxon R. infiata subsp. australis and the depsi- 
done taxon R. fissa, especially as similarities in morphology are also found in some specimens. 
However, as no Australian material was found to contain both depside and depsidone 
chemistry, R. fissa is regarded as a distinct, endemic species. 

Distribution and habitat. The distribution range of R. fissa (Fig. 23) lies within the larger range 
of the two chemical races of R. inflata subsp. australis in the temperate region of Australia. It is 
mostly coastal in occurrence, occupying sections of the coastline in both Tasmania and the 
mainland not usually occupied by R. inflata subsp. australis (divaricatic acid race). In western 
Victoria and South Australia the inland locations of R. fissa are mainly different from the sites 
occupied by R. inflata subsp. australis (Figs 22, 23). 

Variation in morphology appears to be ecophenotypic; in some coastal habitats extreme 
flattening of the thallus occurs, producing a dorsi- ventral form resembling a parmeloid thallus. 
In the dry inland regions of southern Australia the thalli are inflated, small, and very coarsely 
textured. On the islands of Bass Strait and Kangaroo Island, the thalh produce inflated pustules 
or small branchlets on the surface. A lower 'pseudo-cortex' is often produced to cover the 
exposed medulla in some thalli with completely split-open branches. 

Occasionally R. fissa occurs on rocks and produces a much reduced growth form. On Hunter 
Island, Bass Strait, specimens grow on quartzite; Tasmanian collections from Cape Deslacs and 
Bruny Island do not record the rock type. Some of these saxicolous specimens are tiny, single, 
balloon-shaped thalli which closely resemble thalli oiR. inflata subsp. australis (divaricatic acid 
race) collected off fine twigs of Leptospermum sp. in Western Australia. Usually R. fissa is 
corticolous, occurring on a number of different tree species {Acacia spp., Leucopogon par- 
viflora, Leptospermum laevigatum, Pimelea serpyllifolia, and Melaleuca lanceolata); it occurs on 
the mangrove Avicennia marina and has been found growing on fence posts at Mario, Victoria. 
It does not show any substrate preference. 

World distribution. AustraUa. 

Victoria: Linga, 13 km N of, Pink Lakes, 35°06'S, 141°40'E, on dead wood, 1970, GB & JC (H032663); 
Casterton, 51-5 kmN of, 37°20'S, 141°20'E, on Acacia sp., 1973, /?F(MEL1036009); Creswick, 37°26'S, 
143°54'E, on Pinus sp., 1963, /. H. Willis (MEL9335); Lake Burrumbeet, 37°28'S, 143°40'E, on Pinus 
sp., 1945, P. Bibby (MEL10018); Mario, 37°48'S, 148°42'E, on fence post, 1978, JAE (JAE5195); 
Troidea Hill, 37°57'S, 141°19'E, on Banksia sp., 1965, /?F(MEL1026193); Portsea, 38°20'S, 144°42'E, 
on shrub, 1964, /. Williams (MEL1516513); Portland, 38°21'S, 141°36'E, on Mains sp., 1965, RF 
(MEL1516516); Western Port Bay, 38°26'S, 145°08'E, on Avicennia marina, 1978, GNS 
(BRIU2347NS); Wilsons Prom. N of, 39°05'S, 146°25'E, on Acacia sp., 1980, GKS (BRIU3198NS). 
Bass Strait: Hogans I, 39°13'S, 146°59'E, on Leucopogon sp., 1973, JW (MEL10130013); Little Dog I, 



RAMALINA IN AUSTRALIA 



195 





lio 




130 


1 — 

140 






r- 

1S0 






0. 




800km 


^3^ 


^ ^ 




-x 






10- 








f 


. _ _-- 










^ 


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20' 












"\ 


















• 

1 






^' 


r""* 




T ** 


•• 




/ 




30- 














:; 






40- 



Fig. 23 Distribution of R. fissa. 

40°15'S, 148°17'E, on twigs, 1977,/W(MEL1026160); Flinders 1, 40°01'S, 148°02'E, on Leptospermum 
laevigatum, 1969, JW (MEL1019938); Hunter I, 40°31'S, 144°45'E, on quartzite, 1973, T. V. Muir 
(MEL1021245). Tasmania: Table Cape, 40°57'S, 145°43'E, on Acacia melanoxylon, 1970, GB 
(H032296); Eddystone Beach, 40°59'S, 148°19'E, on dead tree, 1973, GB (H032679); Devonport, 
41°11'S, 146°21'E, on Acacia sp., 1980, GNS (BRIU3240NS); Chain of Lagoons R., 41°39'S, 148°18'E, 
on bush, 1973, GB & JC (H032667); Cape Deslacs, 42°57'S, 147°33'E, on rock, 1965, GB & JC 
(H032647); Bruny I, Isthmus Bay, 43°23'S, 147°17'E, on tree, 1982, AA (BRIU3707NS). SoutJi 
Australia: Lock, 25 km WSW of, 33°40'S, 135°20'E, 1967, N. N. Donner (AD97648157); Yararoo 
Gorge, 34°20'S, 138°20'E, 1967, B. Copley (AD97419099); Kangaroo I, 35°40'S, 137°38'E, on Mallee 
bush, 1971, G. Jackson (AD97410372); Coorong, 36°12'S, 139°40'E, 1965, A. C. Beauglehole 
(MEL1020684); Naracoorte, 36°58'S, 140°45'E, on dead wood, 1973, M. Beck (H32559); Eyre 
Peninsula, Cape Catastrophe, 34°40'S, 135°50'E, on Melaleuca sp., 1970, /?F(MEL1020693). 



17. Ramalina nervulosa (Miill. Arg.) des Abb. 

in Bull. Inst. fr. Afr. noire 14: 25 (1952). 

Ramalina farinacea var. nervulosa Miill. Arg. in Flora, Jena 66: 21 (1883). Type: Australia, Queensland, 

Daintree River, Pentzke s.n. (G! - lectotype; divaricatic acid, stenosporic acid, and usnic acid), 
var. nervulosa Plate 11, fig. 3; Plate 14, fig. 7. 

Thallus corticolous, pale green to stramineous, subpendulous to pendulous, usually 4-10 cm 
long, exceptionally to 30 cm; branching sparse with unequal dichotomy producing long 
attenuate branches, often tangled about one another; branch width (0-3-)l-0-l-5(-2-0) mm, 



196 



G. NELL STEVENS 




itiiiimiiiiiimiiiiiiiiiiiiiiiiiiiitfiiiiiiiinniiiiiiiiiiiii iiiiiiiiiiiiiiiiiiiiiiiiiiiiiiitiiiiiiitiiii 



RAMALINA IN AUSTRALIA 197 

branches flat and narrow, when extremely narrow becoming subterete and twisted or undulate 
along the entire length, branches occasionally fuse; cortex matt, marked with longitudinal 
pseudocyphellae which cause the 'nervulose' appearance of this species; several branches arise 
from a well defined holdfast; soralia marginal, round to eUipsoid. Apothecia not seen. 

Chemistry. Divaricatic acid and stenosporic acid and usnic acid. 

Remarks. Miiller [Arg.] (1883) appUed the name R. farinacea var. nervulosa to specimens 
which had been sent to him from Norfolk Island, Australia, Tahiti, the Marianas, and India 
which closely resembled R. farinacea. Des Abbayes (1952) collected and examined sorediate 
specimens from the Ivory Coast (West Africa) and thought they belonged to the same species 
that Miiller had described, but he believed that they were sufficiently distinct from R. farinacea 
to warrant species rank as R. nervulosa (Miill. Arg.) des Abb. (Stevens, 1983fl). 

When the five specimens held at G in the Miiller [Arg.] herbarium were examined, it was 
found that, although their morphology was similar in some instances, each had a quite different 
chemistry and belonged to separate taxa. The majority of the thalli bear the 'nervulose 
markings' (linear pseudocyphellae and longitudinal cracks) which caused the name 'nervulosa' 
to be applied to them; this trait is quite distinctive as remarked upon by des Abbayes. Of the five 
specimens cited by Miiller (1883) only the Australian material from Daintree River Queensland 
contains divaricatic acid and stenosporic acid. This specimen was designated as lectotype by 
Filsoninl971. 

Distribution and habitat. Ramalina nervulosa var. nervulosa occurs along the Queensland 
coastline from latitude 14°-28°S (Fig. 24), and at several inland locations (Stewart River (14°S), 
Barrow's Range (15°S), and Atherton Tableland (17°S)). This taxon is sympatric with its variety 
(var. luciae) for much of its distribution in Australia. The largest specimens collected, which 
exceed 20 cm, grew at Stewart River (14°S) and Turkey (24°S), the reason for such exceptional 
growth is not known. The mangroves Ceriops tagal and Rhizophora stylosa are the usual 
phorophytes for this taxon. 

World distribution: Burma/Bangladesh, Indonesia (Java and Bali), South Africa, and Austra- 
lia. 

Queensland: Stewart R., 14°05'S, 143°38'E, 1900, Johnston (MEL 4443); Bartons Range, 15°36'S, 
145°04'E, on shrub, 1983, M. Godwin (C2436); Mossman Rifle Range, 16°20'S, 145°45'E, on Ceriops 
tagal, 1983, GNS (BRIU4128NS); Trinity Inlet (Cairns); 16°56'S, 145°46'E, on Ceriops tagal, 1976, D. 
Tarte (BRIU2589NS); Hinchinbrook I, 18°17'S, 146°13'E, on Ceriops tagal, 1979, GNS (BRIU3131NS); 
Eimeo, 21°09'S, 149°09'E, on Rhizophora sp., 1976, GNS (BRIU3544NS); St Lawrence, 22°21'S, 
149°32'E, on Excoecaria agallocha, 1976, NCS (BRIU2131NS); Port Alma, 23°35'S, 150°51'E, on 
Rhizophora stylosa, 1975, RWR (BRIU698RR); Gladstone, 23°50'S, 151°16'E, on Rhizophora stylosa, 
1980, GNS (BRIU3348NS); Rodds Peninsula, 24°02'S, 151°40'E, on Ceriops tagal, 1975, GNS 
(BRIU1292NS); South Stradbroke I, 27°55'S, 153°30'E, on Ceriops tagal, 1975, H. T. Clifford 
(BRIU76RR). 

var. dumeticola (Krog & Swinscow) N. Stevens Plate 11, fig. 2. 

in Lichenologist 15: 228 (1983). 

Ramalina dumeticola Krog & Swinscow in Norw. J. Bot. 23: 163 (1976). Type: Kenya, Coast Province, 
Kwale District, Krog 3K30/127 (O! - holotype, BM! - isotype; homosekikaic acid (M), sekikaic acid 
(M), 4'-0-methylnorhomosekikaic acid (M), ramalinolic (t), 4'-0-demethylsekikaic acid (t), 4'-0- 
methylnorsekikaic acid (t) and usnic acid). 

Thallus corticolous, pale green to yellow-green, tufty, subpendulous, 4-6 cm long; branching 
dichotomous, moderate to dense but never intricately branched; branch width (0-2)0-5-l-0(- 

Plate 11 Thallus growth forms in the Australian Ramalinae. Scale in mm. Fig. 1 R. pacifica. Fig. 2 R. 
nervulosa var. dumeticola. Fig. 3 R. nervulosa var. nervulosa. Fig. 4 R. nervulosa var. luciae. Fig. 5 R. 
subfraxinea var. confirmata. Fig. 6 R. subfraxinea var. leiodea. Fig. 7 R. subfraxinea var. subfraxinea 
(narrow form). Fig. 8 R. subfraxinea var. norstictica (type specimen). 



198 



G. NELL STEVENS 





120 




130 


140 




150 









800km 


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- 










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Fig. 24 Distribution of R. nervulosa var. nervulosa. 



2-0) mm, branches compressed, flat, usually canaliculate for part of their length, some branches 
subterete distally, apices attenuate, forked or broken off and appearing blunt; surface matt or 
shiny, smooth, faint pseudocyphellae sometimes present; holdfast deUmited; soralia marginal, 
round, ellipsoid or elongate. Apothecia not seen. 

Chemistry. Homosekikaic acid (M), sekikaic acid (M), 4'-0-methylnorhomosekikaic acid (m), 
4'-0-demethylsekikaic acid (t), ramalinolic acid (t), 4'-0-methylnorsekikaic acid (t) [the last two 
acids are rarely detected with tic] and usnic acid. 

Remarks. Initially the Australian material of this taxon was thought to be a morphotype of R. 
peruviana as the two are sympatric in occurrence. However, the shape of the soralia and 
branching pattern of the two sets them apart. The morphology of R. nervulosa var. dumeticola 
closely resembles that of R. farinacea. Although both R. peruviana and R. nervulosa var. 
dumeticola contain acids in the sekikaic aggregate, the presence of 4'-0-methylnorhomosekikaic 
acid as a minor metabolite in the latter taxon and not in the former, and the constant occurrence 
of ramalinolic acid in R. peruviana but not in R. nervulosa var. dumeticola distinguishes the two 
on chemical grounds. 

Distribution and habitat. The occurrence of jR. nervulosa var. dumeticola in Australia at 
subtropical latitudes appears to be an anomaly (Fig. 25). Previous research (Stevens, 1983fl) has 
shown its distribution to be mostly equatorial and northern hemisphere tropical (to latitude 
15°N), penetrating to latitude 8°S (based on known collections). Its presence in the mountains of 



RAMALINA IN AUSTRALIA 



199 




Fig. 25 Distribution of R. nervulosa var. luciae • and var. dumeticola *. 



New Guinea at latitude 6°S (established during this present research) is in accord with this 
distribution pattern. The subtropical occurrence of this taxon in Australia in the mountain areas 
between latitudes 26°S-30°S cannot be explained, nor can the complete absence of this taxon 
from any coastal habitats. 

World distribution. India, Indonesia (Java), Philippines, Marianas Is, Kenya, Ivory Coast, 
Principe, Ascension, and Australia. 

Queensland: Bunya Mts, 26°50'S, 151°40'E, on understorey shrub, 1979, GNS (BRIU2971NS); Mt Castle, 
2 km SW of, 27°58'S, 152°23'E, on rainforest twig, 1983, NCS (BRIU3677NS); Cunninghams Gap, Mt 
Cordeaux track, 28°03'S, 152°23'E, on rain-forest twig, 1979, GNS (BRIU3697NS); The Head, Moss 
Gardens, 28°15'S, 152°30'E, on twigs of rain-forest shrub, 1981, GNS (BRIU3622NS); O'Reillys, 
Botanical Walk, 28°15'S, 153°08'E, on rain-forest shrub, 1983, GNS (BRIU3731NS); Gamboobal State 
Forest, 28°14'S, 152°20'E, on dead tree on mountain top, 1983, GNS (BRIU3737NS); Queen Marys Falls, 
5 km N of, 28°20'S, 152°22'E, on rain-forest tree, 1983, GNS (BRIU3736NS). New South Wales: 
ChaelundiMt,37kmNofEbor,30°04'S, ISTll'E, Acacia orrites, 1978, D. Verdon{CBGlS09394); Cattle 
Ck State Forest, 12 km NNE of Dorrigo, 30°15'S, 152°03'E, on Banksia integrifolia, 1978, D. Verdon 
(CBG8203532). 



var. /uc/ae (Molho et al.) N. Stevens 



Plate 11, fig. 4. 



in Lichenologist 15: 227 (1983). 

Ramalina luciae Molho, Bodo, Culb. & C. Culb. in Bryologist 84: 396 (1981). Type: Fiji, Kambara (= 



200 G. NELL STEVENS 

Kabara) Island, Salvat B. 358a (DUKE - holotype; 'sekikaic, homosekikaic, 4'-0-demethylsekikaic and 
4'-0-methylnorsekikaic acids and usnic acid'). 

Thallus corticolous, pale green to stramineous, subpendulous to pendulous, up to 7 cm long, 
exceptionally to 18 cm; branching sparsely dichotomous, producing extreme elongation 
between dichotomies; branch width (0-3)0-5-l-0(-2-0) mm, branches flattened at the base 
but subterete to terete towards the apices, slightly twisted along the whole length, fusion 
between branches common, causing a tangled growth form; cortex matt to shiny; longitudinal 
pseudocyphellae usually present but rarely becoming laminal soralia; simple cracks in the 
cortex and splitting along the margin occurs; several basal branches arise from a well defined 
holdfast. 

Apothecia rare, lateral, disc 2 mm diam, concave to ellipsoid to fusiform, sometimes 
appearing 3,4-celled; straight or rarely slightly curved (10-)12-16(-18) x 4-5(-6) /tm. 

Chemistry. Sekikaic acid (M), 4'-0-methylnorsekikaic acid (m), 4'-0-demethylsekikaic acid (m 
or t), homosekikaic acid (t), ramalinoUc acid (t), 4'-0-methylnorhomosekikaic acid (t). 

Remarks. Ramalina nervulosa var. luciae was described by Molho, Bodo, Culberson & C. 
Culberson (1981) from a specimen collected in Fiji. Similar material held at G as a syntype of/?. 
farinacea var. nervulosa Miill. Arg. , was collected from Tahiti last century by Jardin. This taxon 
is a common maritime Ramalina along the eastern coastline of tropical Australia. 

Because of the similarity in morphology between R. nervulosa var. nervulosa and R. nervulosa 
var. luciae, chemical differences had to be used as the main criteria to separate them. The rank of 
variety was given to each taxon in accordance with the guidelines set out by Hawksworth (1976). 
Apart from chemical difference, their distribution range when considered on a global scale 
shows two distinct patterns (Stevens, 1983fl). 

The chemistry of two sorediate Ramalina species resembles that of R. nervulosa var. luciae 
(viz. R. peruviana Ach. and R. intermediella Vainio) but both of these have a distinct 
morphology which distinguishes them from the morphology oiR. nervulosa var. luciae. 

Distribution and habitat. The distribution of R. nervulosa var. luciae (Fig. 25) covers a coastal 
region from approximately latitudes 15°-27°S in Queensland. For most of its range this taxon is 
sympatric with both R. nervulosa var. nervulosa and R. pacifica. 

The majority of the collections was made at sea-level from mangrove communities, the main 
phorophytes being Ceriops tagal, and to a lesser extent Avicennia marina and Rhizophora 
stylosa which grow on the sheltered, landward fringe of the mangroves. The most northerly 
collections in Austraha are from inland sites, e.g. Butchers Hill, Lakeland, latitude 15°52'S, 
and Atherton Tableland, latitude 17°16'S. Both of these sites are approximately 45 km from 
the coast at an altitude of 900 m, indicating cooler habitat requirements at lower latitudes; 
a tendency observed in the other members of the tropical R. farinacea complex (Stevens, 
1983fl). 

At latitude 21°S (Eimeo Beach, Mackay) this species grows prolifically on the landward fringe 
of Rhizophora stylosa communities; in this habitat fertile specimens as well as very long 
specimens occur. 

World distribution. Sri Lanka, Indonesia (Celebes), Kenya, Australia, New Hebrides, Fiji, 
Cook Is, and Tahiti. 

Queensland: Lakeland, Butchers Hill, 15°52'S, 144°55'E, on rain-forest shrub, 1983, GNS 
(BRIU4108NS); Mossman, 16°20'S, 145°24'E, on Pinus sp. in garden, 1983, C. McCracken 
(BRIU4241NS); Yungaburra Rd Atherton, 17°16'S, 145°29'E, on Casuarina sp., 1983, HS 
(CBG8302450); Hinchinbrook 1, 18°17'S, 146°13'E, on Ceriops tagal, 1979, GNS (BR1U3137NS); Eimeo, 
21°09'S, U9°09"E, on Rhizophora stylosa, 1976, GA^5(BRIU3542NS); St Lawrence, 22°21'S, 149°32'E,on 
Ceriops tagal, 1980, GNS (BR1U3600NS); Corio Bay, 22°58'S, 150°46'E, on Ceriops tagal, 1974, D. Tarte 
(BRIU1168NS); Gladstone, 23°50'S, 151°16'E, on Ceriops tagal, 1980, GNS (BRIU3566NS); Tannum 
Sands, 23°57'S, 151°16'E, on Ceriops tagal, 1975, RWR (BRIU833RR); Turkey, 24°06'S, 151°40'E, on 
Rhizophora stylosa, 1975, GA^5(BRIU280NS);Toogoom,25°15'S, \52°m'^, on Ceriops tagal, 1983, GNS 
(BRIU4149NS); Serpentine Ck, 27°23'S, 151°39'E, on Ceriops tagal, 1975, GNS (BRIU1044NS). 



RAMALINA IN AUSTRALIA 201 

18. Ramalina pacWca Asah. Plate 11, fig. 1; Plate 14, fig. 8. 

in /. Jap. Bot. 15: 213 (1939). Type: Micronesia, Marianas, Saipan Island, 1925, Kimura s.n. (TNS! - 

lectotype; salazinic acid and usnic acid). 
Ramalina insularum Magnusson in Ark Bot. 3 (10): 361 (1956). Type: Fiji Is., 1862, Daemel s.n. (H! - 

lectotype; salazinic acid and usnic acid). 

Thallus corticolous, rarely saxicolous, grey-green, tufty, subpendulous to pendulous, 4-8 cm 
long, exceptionally to 32 cm; branching dichotomous, sparse to moderately dense but extremely 
variable; branch width (0-5-)l-4 mm, branches compressed, flat, some splitting along the 
margins; broad in the basal region but gradually narrowing towards the apices or narrow in the 
basal region and continuing the same width to the apices; apices attenuate, mostly forked; cortex 
matt or shiny, smooth or with linear pseudocyphellae slightly or strongly developed; holdfast 
delimited; soralia marginal and laminal, round to ellipsoid (Plate 1, fig. 3), often coalescing, the 
edges re volute. 

Apothecia rare, marginal, disc 1-3 mm diam., concave, plane to convex, with edges 
sometimes incised at maturity; spores fusiform or ellipsoid, (Plate 4, fig. 3), straight or very 
slightly curved, 12-16(-20) x 4-6 /am. 

Chemistry. Salazinic acid ± protocetraric acid and usnic acid. 

Remarks. The name Ramalina pacifica was given to sorediate material found on the Saipan 
Islands (Marianas) and on Hachijo Island (Japan) by Asahina (1939); he stated that lichenol- 
ogists had identified his species as various varieties of R. farinacea. Asahina also commented 
that some thalh of R. pacifica contained both salazinic and sekikaic acids. It appears that the 
sekikaic acid he observed could have come from a contaminant, perhaps R. intermediella or R. 
nervulosa var. dumeticola, as both of these species are maritime and grow in association with R. 
pacifica. 

Stevens (1983a) placed R. pacifica in the tropical-subtropical group of the R. farinacea 
complex. It is the only taxon in this group which has a depsidone chemistry. In ancient times it 
may have been contiguous with the protocetraric acid species of R. farinacea found on 
Macquarie I. (latitude 55°S), but the links are now broken except for the presence of trace 
amounts of protocetraric acid in some specimens of R. pacifica. 

Ramalina insularum Magnusson, a synonym of R. pacifica, was collected from Fiji. Its 
morphology more closely resembles that of R. nervulosa var. luciae material collected from Fiji 
than it does the Australian R. pacifica, which illustrates the phenomenon of convergent 
morphology of these two Fijian specimens which are sympatric. 

Distribution and habitat. The distribution range of R. pacifica covers an area which is mainly 
subtropical to warm temperate (Fig. 26), although several collections have been made in tropical 
Austraha, e.g. Hinchinbrook I. (latitude 18°S) and Shute Harbour (latitude 20°S). In these areas 
the rainfall exceeds 2000 mm p. a. and the thalli are very narrow and delicate with smooth, shiny 
surfaces, without pseudocyphellae. Collections from the drier latitudes along the Queensland 
coast (latitudes 22°-24°S which receive 800-1100 mm p. a.) comprise thalli which produce 
numerous pseudocyphellae along the whole length of the branches; these thalh are generally 
broader than the thalh from the higher rainfall habitats. Only one inland collection of/?, pacifica 
was made, at Atherton Tableland. 

The main phorophytes for this taxon are the mangroves Ceriops tagal and Avicennia marina. 
Ramalina pacifica attains its maximum growth at latitudes 27°-28°S (Moreton Bay), annual 
rainfall 1100-1500 mm p. a. In two relatively undisturbed well-established communities of 
Avicennia marina some exceptional specimens 32 cm long with branches 2-3 mm wide were 
found. Fertile specimens also occurred in this environment. The majority of the thalli produced 
pseudocyphellae as well as both marginal and laminal soralia. In areas of Moreton Bay where R. 
pacifica grew on Ceriops tagal, the thalli were small (up to 5 cm long) with narrow branches (to 
1-0 mm wide), and had a smooth cortex which lacked pseudocyphellae or produced them at the 
base of the branches only. 

From latitude 28°S to the limit of R. pacifica distribution at 35°S, this taxon occurs on 



202 



G. NELL STEVENS 




Fig. 26 Distribution of R. paciftca. 



Avicennia marina {Ceriops tagal does not extend beyond latitude 28°S). Pseudocyphellae are 
usually present on the branches which reach 8 cm in length with a maximum width of 2 mm. The 
annual rainfall in this section of coastline is between 1000-1500 mm p. a. 

It was not able to be established whether the amount of rainfall or the type of substrate had the 
greater influence on the variation in morphology of this taxon. 

Ramalina pacifica is rarely saxicolous but occurs on rock at Norries Head, New South Wales, 
in association with R. subfraxinea var. confirmata, another taxon which is usually corticolous. 
Presumably the rocks in this area of coastline are able to produce a microhabitat which is 
equivalent to that found in tree communities on other parts of the coast, enabling both taxa to 
become estabhshed on the cliff face. 

World distribution. Burma/Bangladesh, Japan, Indonesia (Java), Marianas Is, South Africa, 
AustraUa, New Hebrides, New Zealand, and Cook Is. 

Queensland: Tinaroo Dam, 17°15'S, 145°36'E, on Plumeria sp. , 1983, GNS (BRIU41NS); Hinchinbrook I, 
18°17'S, 146°13'E, on Ceriops tagal, 1979, GNS (BRIU3441NS); Bowen, 19°59'S, 148°22'E, on Lumnit- 
zera racemosa, 1979, GA^5 (BRIU3284NS); Shute Harbour, 20°16'S, 148°47'E, on Ceriops tagal, 1979, 
GN5(BRIU3283NS); Brampton 1, 20°49'S, 149°18'E,/. M. G//fcer^(H032317);Eimeo,21°09'S, 149°09'E, 
on Rhizophora stylosa, 1976, GNS (BRIU1369NS); St Lawrence, 22°21'S, 149°32'E, on Excoecaria 
agallocha, 1976, GNS (BRIU2127NS); Keppel Sands, 23°21'S, 150°47'E, on Excoecaria agallocha, 1975, 
RWR (BRIU779RR); Port Alma, 23°35'S, 150°51'E, on Rhizophora sp., 1975, RWR (BRIU699RR); 
Gladstone, 23°50'S, 151°16'E, on Rhizophora stylosa, 1980, GNS (BRIU3570NS); Tannum Sands, 



RAMALINA IN AUSTRALIA 203 

23°57'S, 15ri6'E, on Rhizophora stylosa, 1980, GNS (BRIU3599NS); Turkey, 24°06'S, 151°37'E, on 
Rhizophora stylosa, 1975, GNS (BRIU1266NS); Rodds Peninsula, 24°02'S, 15r40'E, on Ceriops tagal, 
1975, GNS (BRIU1153NS); Round Hill Hd., 24°10'S, 151°54'E, on Rhizophora stylosa, 1975, RWR 
(BRIU915RR); Eraser I, Wathumba Ck, 24°59'S, 152°55'E, on Rhizophora stylosa, 1975, GNS 
(BRIU3597NS); Burrum Hds, 25°10'S, 152°35'E, on Rhizophora stylosa, 1975, GNS (BRIU1129NS); 
Urangan Beach, 25°19'S, 152°55'E, on Callitris columellaris , 1978, GNS (BRIU2841NS); Tin Can Bay, 
25°55'S, 153°00'E, GB (H032319); Noosa Hds, 26°24'S, 152°07'E, on Rhizophora stylosa, 1975, GNS 
(BRIU1807NS); Deception Bay, 27°10'S, 153°02'E, on Avicennia marina, 1978, GNS (BRIU2561NS); 
Tingalpa Ck, 27°28'S, 153°11'E, on Avicennia marina 1977, GNS (BRIU1756NS); Coochiemudlo I, 
27°35'S, 153°20'E, on Rhizophora stylosa, 1977, GN5(BRIU1654NS); Little Rocky Pt,27°38'S, 153°18'E, 
on Ceriops tagal, 1977, GNS (BRIU1438NS); Stradbroke I, Swan Bay, 27°45'S, 153°26'E, on Ceriops 
tagal, H. T. Clifford (BRIU3075RR). New South Wales: Brunswick Hds, 28°32'S, 153°32'E, on Callitris 
sp., 1980, GNS (BRIU3486NS); Yamba, 29°27'S, 153°20'E, on Avicennia marina, 1977, GNS 
(BRIU2220NS); Boambee Ck, 30°22'S, 153°05'E, on Avicennia marina, 1977, GNS (BRIU2032NS); 
Nambucca Hds, 30°39'S, 153°00'E, on Avicennia marina, 1977, GNS (BRIU2001NS); Port Macquarie, 
31°27'S, 152°54'E, on Avicennia marina, 1977, GNS (BRIU1989NS); Patonga, 33°30'S, 151°20'E, on 
Avicennia marina, 1978, GNS (BRIU2308NS); Towra Pt., 34°01'S, 151°13'E, on Avicennia marina, 1977, 
C. Scarlett (BRIU634CS); Sussex Inlet, 35°19'S, 150°36'E, on Avicennia marina, 1977, RWR 
(BRIU2226NS). 

19. Ramalina subfraxineaNyl. 

in Bull. Soc. Linn. Normandie II, 4: 139 (1870). Type: South America, Columbia, Turbaco, Nova 
Granata, 1844, Goudot s.n. (PC! - lectotype; H! - isotype; sekikaic acid, homosekikaic acid, an 
unknown and usnic acid). 

var. subfraxinea Plate 11, fig. 7; Plate 12, figs 3, 7; Plate 15, fig. 2. 

Thallus corticolous, pale green to grey-green, erect to subpendulous, up to 3 cm long, 
exceptionally to 12 cm; branching subdichotomous, sparse; branch width 0-5-1 •0(-3-5) mm, 
branches compressed, narrow and canaliculate with attenuate apices or broad and flat with blunt 
apices; surface matt or shiny, marginal tuberculate pseudocyphellae frequent, laminal punc- 
tiform pseudocyphellae on broad branches; holdfast delimited; soralia absent. 

Apothecia sparse to numerous, marginal and subterminal; disc 1-4 mm diam. , concave, plane 
to convex; margin entire; spores ellipsoid, straight to slightly curved, 11-12(-16) x 4-5(-6)/xm. 

Chemistry (Australian taxa). Sekikaic acid, homosekikaic acid, ramalinolic acid, 4'-0- 
demethylsekikaic acid, paludosic acid, 4'-0-methylnorsekikaic acid, and usnic acid or divaricatic 
acid ± faint sekikaic acid and usnic acid. 

Remarks. Nylander (1870) reported material of R. subfraxinea from Nova Granata (Goudot), 
Mauritius (Cretan ex hb. Lenorm.), Bourbon I. (Boivin), and the Marianas Islands 
(Gaudichaud). Examination and chemical analysis of material held at H-NYL and PC revealed 
that the material from Nova Granata contained either the acids of the sekikaic aggregate or 
divaricatic acid, whereas all of the material from Mauritius, Bourbon I., and the Marianas Is. 
contained boninic acid. Those collections found to contain boninic acid belong to the species R. 
leiodea (Stevens, 1982), regarded as R. subfraxinea var. leiodea in the present study. 

Material from Nova Granata, collected by Goudot, is held at both PC and H; the material 
from PC comprises several thalli in good condition on a twig, whereas the specimen held at H is 
small. Every thallus on the twig held at PC was analysed and the thallus herein designated as 
lectotype contains sekikaic acid, homosekikaic acid, and faint 4'-0-demethylsekikaic acid. The 
morphology of this thallus fits the description in the protologue: 'subsimilis Ramalinae fraxineae 
minori et attenuatae, sed sporis (subcurvulis) rectioribus tenuioribusque (longit 0,011-18 
millim. crassit 0,0035-0,0045 millim.)'. 

It was found that several thalli on the twig contain divaricatic acid, although morphologically 
identical to those containing sekikaic acid. The thallus containing sekikaic acid was chosen for 
lectotypification in preference to a thallus containing divaricatic acid because of the large 
number of sekikaic acid specimens found in Australia compared to the few specimens which 



204 



G. NELL STEVENS 



contain divaricatic acid. Another reason for this choice was the record (Krog & Swinscow, 1976) 
of the presence of the sekikaic acid aggregate in both PC and H collections of R. subfraxinea. 

Distribution and habitat. Ramalina subfraxinea var. subfraxinea is tropical in distribution, 
reaching its southern limit at latitude 24°S, on the eastern coast of Australia; on the northern 
coastline it has been collected from three areas (Darwin, Gunn Pt, and Gove) at latitude 12°S 
(Fig. 27). This taxon has two chemical races (a) a sekikaic acid aggregate race and (b) a 
divaricatic acid race. The former is widely distributed from 12°S-24°S, whereas the divaricatic 
acid race has a restricted range between latitudes 15°5rS and 18°17'S (Fig. 31), where both races 
are sympatric. 

The occurrence of R. subfraxinea var. subfraxinea is almost entirely coastal, with only one 
inland collection from Butchers Hill (15°51'S) where both races occur on shrubs in a low 
closed-format community. Thalli from this area contain divaricatic acid + trace amounts of 
sekikaic acid or sekikaic acid only indicating a close affinity between the two acid races and 
strengthening the argument for not separating each acid as a different variety of R. subfraxinea. 

The main phorophytes for this taxon are Ceriops tagal, Rhizophora stylosa, Lumnitzera 
racemosa, Excoecaria agallocha, and several rain-forest trees which occur along the tropical 
beaches. 

World distribution. India?, Australia, and South America (Colombia). 

Northern Territory: Darwin, Stuart Park, 12°27'S, 130°50'E, on Ceriops tagal, 1981 , GNS (BRIU3496NS); 
Gunn Point, 12°08'S, 130°59'E, on Lumnitzera racemosa, 1979, E. Hegerl (BRIU3115NS); Gove Airport, 





120 




ido 




^io 






160 







SOOkm 


^^^ 


-^ 




f\ 




10- 








vi' 


\ 


. / 


i 


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/ 






\ / 




V 




/ 














> 


i 20- 




' " 














'\ 


X 
















I 


. \ 


L 


/" ol 


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^y^ 

D 


^ 

% 


j 30- 






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" 






^ 


■L 


40- 



Fig. 27 Distribution of R. subfraxinea var. subfraxinea. 



RAMALINA IN AUSTRALIA 205 

12°18'S, 136°50'E, on Croton sp., 1982, N. Sammy 820576. Queensland: Princess Charlotte Bay, Marrett 
R., 14°25'S, 144°09'E, on Ceriops tagal, 1979,7. Davie (BRIU2973NS); Lakeland, Butchers Hill, 15°51'S, 
144°53'E, on rain-forest shrub, 1983, G.N. (BRIU4107NS); Daintree R. (south branch), 16°16'S, 
145°25'E, on Ceriops tagal, 1983, GNS (BRIU4136NS); Newell Beach, S Saltwater Ck 16°25'S, 145°25'E, 
on Ceriops tagal, 1983, C. McCracken (BRIU4234NS); Mossman Rifle Range, 16°27'S, 145°22'E, on 
Ceriops tagal, 1983, GNS (BRIU4126NS); Trinity Inlet, 16°56'S, 145°45'E, on Ceriops tagal, 1975, D. 
Tarte (BRIU2464NS); North Mission Beach, 17°52'S, 146°06'E, on Rhizophora sp., 1979, NCS 
(BRIU2663NS); South Mission Beach, 17°55'S, 146°06'E, on Ficus sp., 1983, GNS (BRIU4161NS); 
Hinchinbrook I, The Haven, 18°18'S, 146°10'E, on Lumnitzera sp., 1979, GNS (BRIU3469NS); Towns- 
ville, 19°15'S, 146°50'E, on Ceriops tagal, 1975, B. Bailment (BRIU3119NS); Shute Harbour, 20°15'S, 
148°47'E, on rain-forest tree, 1979, GNS (BRIU3139NS); Hallidays Bay, 20°56'S, 149°01'E, on Ceriops 
tagal, 1983, GNS (BRIU4122NS); Eimeo, 21°09'S, 149°10'E, on Rhizophora stylosa, 1976, GNS 
(BRIU1353A.NS); St Lawrence, 22°21'S, 149°32'E, on Excoecaria agallocha, 1976, GNS (BRIU2132NS); 
Corio Bay, 22°58'S, 150°45'E, on Rhizophora sp., 1975, D. Tarte (BRIU1169NS); Rodds Peninsula, 
24°02'S, 15r40'E, on Rhizophora stylosa, 1975, GNS (BRIU1278NS); Turkey, 24°06'S, 151°37'E, on 
Rhizophora stylosa, 1975, GNS (BRIU1150NS). 



var. conRrmata (Nyl.) N. Stevens, comb. nov. 

Plate 11, fig. 5; Plate 12, figs 4, 10; Plate 15, figs 6, 8. 

Ramalina fraxinea [subsp.] confirmata Nyl. in Bull. Soc. linn. Normandie II, 4: 138 (1870). Type: 
Australia, Swan R., 1846, Verreaux s.n. (H-NYL 37423! - lectotype; cryptochlorophaeic acid and usnic 
acid). 

Thallus corticolous, rarely saxicolous, grey-green, rigid, erect to subpendulous, up to 4 cm 
long, exceptionally to 11 cm; branching subdichotomous, sparse to moderate; branch width 
(0-5-)2-4(-10) mm, branches compressed, narrow and canaliculate to broad and flat, apices 
attenuate or blunt; surface smooth to rugose, short linear pseudocyphellae along branches or 
basally only; holdfast delimited; soralia absent. 

Apothecia common, marginal and subterminal, rarely laminal, small thalli produce subter- 
minal apothecia subtended by a spur; disc 2-3 mm diam., concave or plane; margin entire, 
slightly crenate, often incised at maturity; spores ellipsoid, straight or curved (8-)10-12(-14) x 
4-5 /xm. 

Chemistry. Cryptochlorophaeic acid (M) ± paludosic acid (m), 4'-0-methylcryptochlorophaeic 
acid (m) and usnic acid. 

Remarks. The name R. fraxinea subsp. confirmata Nyl. was given to material sent to Nylander 
by Verreaux, supposedly from Swan River, Western Australia. However, it is doubtful whether 
the specimens came from that area as Verreaux did not personally collect in Western Australia 
(Lamy, in lift.) and no further specimens have been found there. Material held at H-NYL, 
numbers 37421, 37423, all contain cryptochlorophaeic acid. 

Until the present study both boninic acid and cryptochlorophaeic acid taxa were included in 
R. leiodea (Stevens, 1982); however, the policy adopted herein gives varietal status to different 
chemical races within a complex if the distribution patterns differ from one another. The boninic 
acid race therefore retains the name R. subfraxinea var. leiodea and the cryptochlorophaeic acid 
race has been give the name R. subfraxinea var. confirmata. 

Distribution and habitat. This taxon is the most common maritime Ramalina along the eastern 
coastline of Australia, occurring from latitude 10°S (Sue L , north of Cape York) to latitude 3rS 
(Fig. 28). 

Although mostly found along the shoreline, R. subfraxinea var. confirmata has been collected 
from the coastal ranges at Montville, Kenil worth, Bartle Frere, and Atherton Tableland. 

In the mangrove communities it grows on Avicennia marina and Rhizophora stylosa. It has 
also been collected on many coastal shrubs and on Casuarina equisetifolia and Callitris 
columellaris , indicating it has no substrate specificity. This taxon was collected off a rock 
substrate at Norries Head, New South Wales, where metamorphic rocks form a sheer cliff facing 



206 



G. NELL STEVENS 




Fig. 28 Distribution of R. subfraxinea var. confirmata • and var. leiodea O. 

the ocean. Such exposure is comparable with the seaward fringe of the mangroves, the position 
usually occupied by this taxon. 

On a global scale the distribution pattern of the cryptochlorophaeic acid taxon differs from 
that of the boninic acid taxon. Ramalina subfraxinea var. confirmata appears to extend 
eastwards from Australia only as far as New Caledonia, but occurs to the north and in the Indiaa 
Ocean, whereas R. subfraxinea var. leiodea is common on most of the islands in the Pacific 
Ocean, as well as occurring in the Indian Ocean region. 

World distribution. Timor, Zanzibar, Australia, and New Caledonia. 

Queensland: Sue Island, 10°12'S, 142°49'E, on dune shrub, 1981, J. Clarkson 3957; Ingram I, 14°26'S, 
144°50'E, on Surina maritima, 1984, M. Godwin (BRIU4282NS); Turtle Group 1, 14°42'S, 145°11'E, on 
beach shrub, 1979, /. Davie (BRIU3145NS); Cooktown, 15°28'S, 145°15'E, on Rhizophora sp., 1983, 
G.N. (BRIU4112NS); Green Island, 16°45'S, 145°59'E, on beach shrub, 1976, JAE (JAE 2593); Trinity 
Inlet, 16°56'S, 145°46'E, on Ceriops tagal, 1976, D. Tarte (BRIU2252NS); North Mission Beach, 17°52'S, 
146°06'E, on Rhizophora sp., 1979, NCS (BRIU2663NS); Hinchinbrook I, 18°14'S, 146°13'E, on 
Lumnitzerasp., 1979, GA^5(BRIU3465NS);Townsville, 19°15'S, U6°50'E, on Rhizophora sty losa, 1975, 
B. Bailment (BRIU3117NS); Bowen, Dry Ck Rd, 20°04'S, 148°22'E, on Rhizophora apiculata, 1979, GNS 
(BRIU3133NS); Eimeo, Sunset Beach, 21°09'S, 149°09'E, on Rhizophora sp., 1976, GNS 
(BRIU1353NS); Keppel Sands, 23°21'S, 150°47'E, on Avicennia marina, 1975, RWR (BRIU900RR); Port 
Alma, 23°35'S, 150°51'E, on Rhizophora sp., 1975, RWR (BRIU695RR); Gladstone, 23°50'S, 15ri6'E, 
on Rhizophora stylosa, 1980, GNS (BRIU3456NS); Turkey, 24°06'S, 151°37'E, on Ceriops tagal, 1975, 



RAMALINA IN AUSTRALIA 207 

GNS (BRIU1271); Round Hill Head, 24°10'S, 151°54'E, on Rhizophora stylosa, 1975, RWR 
(BRIU916RR); Burnett Hds, 24°45'S, 152°25'E, on Casuarina sp. , 1975, GNS (BRIU4296); Burrum Hds, 
25°10'S, 152°35'E, on Rhizophora stylosa, 1975, GNS (BRIU1121NS); Urangan, 25°19'S, 152°55'E, on 
Callitris columellaris , 1978, GNS (BRIU2840NS); Kin Kin Ck, 26°13'S, 153°00'E, on Avicennia marina, 
1979, GNS (BRIU3214NS); Maroochy R, 26°42'S, 153°05'E, on Avicennia marina, 1976, GNS 
(BRIU1335NS);BribieI,27°02'S, \5yQWE,onLumnitzeraracemosa, 1978, GA^5(BRIU2283NS); Whyte 
I, 27°23'S, 153°10'E, on Avicennia marina, 1978, GNS (BRIU2594NS); Mud I, 27°20'S, 153°15'E, on 
Avicennia marina, 1978, GNS (BRIU2760NS); Coocbiemudlo I, 27°32'S, 153°17'E, on Callitris columel- 
laris, 1975, GNS {IV.lllll); Bilinga, 28°08'S, 153°31'E, on dead tree, \9^\,AA (BRIU4276NS); Norries 
Hd, 28°18'S, 153°34'E, on metamorphic rock, 1980, GNS (BRIU3665NS); Byron Bay, 28°39'S, 153°37'E, 
on dune shrub, 1978,/?W/?(BRIU3883*NS); Yamba,29°27'S, 153°2Q"E, on Avicennia marina, 1977, GNS 
(BRIU3177NS); Port Macquarie, 3r27'S, 152°54'E, on Ficus sp., 1975, M£ (JAE 1089); Hunter R., Ash 
I, 32°56'S, 151°46'E, 1845,/. P. Verreaux (PC). 

var. leiodea (Nyl.) N. Stevens, comb. nov. Plate 11, fig. 6; Plate 12, fig. 8; Plate 15, fig. 3. 

Ramalina subfraxinea [subsp.] leiodea Nyl. in Bull. Soc. linn. Normandie II, 4: 141 (1870). - Ramalina 

leiodea (Nyl.) Nyl., Lich. Nov. Zel. : 22 (1888). Type: New Caledonia, 'Donne par le Musee Colonial', 

1861, Deplanche s.n. (PC! - lectotype; boninic acid aggregate). 
Ramalina boninensis Asah. in/. Jap. Bot. 14: 253 (1938). Type: Bonin Islands, Ogasawara I., Hahajima, 

1936, Kusaka s.n. (TNS! - holotype; boninic acid aggregate). 
Ramalina boninensis f. subcalicariformis Asah. in /. Jap. Bot. 15: 211 (1939). Type: Bonin Islands, 

Ogasawara I., Hahajima, 1936, Kusaka s.n. (TNS! - holotype). 

Thallus corticolous, grey-green, caespitose, rigid, erect to subpendulous, up to 4 cm high, 
exceptionally to 9 cm; branching subdichotomous, sparse to moderate; branch width (0-5-)2-4 
mm, branches compressed, flat or canaliculate when narrow, apices attenuate; surface matt, 
smooth to rugose, short linear pseudocyphellae sometimes present; holdfast deUmited; soralia 
absent. 

Apothecia common, marginal and subterminal, rarely laminal, small thalli produce subter- 
minal apothecia subtended by a spur; disc 2-3 mm diam., concave to plane; margin entire or 
incised at maturity; spores eUipsoid, straight or slightly curved; 10-12(-16) x 4-5 /xm. 

Chemistry. Boninic acid (M), 2-0-methylsekikaic acid (m), 2,4'-di-0-methylnorsekikaic acid 
(m), 4'-0-methylpaludosic acid (m), 4,4'-di-0-methylcryptochlorophaeic acid (m), and usnic 
acid. 

Remarks. Prior to the present study R. subfraxinea var. leiodea had species status. The history 
of this taxon, however, is one of name changing since its first identification by Nylander in 1859. 
Specimens of this taxon have been identified as follows: Ramalina Ecklonii Sprengel (Nylander, 
1859); Ramalina calicaris f. Ecklonii (Sprengel) Nyl. (Nylander, 1861); Ramalina subfraxinea 
[subsp.] leiodea Nyl. (Nylander, 1870) and Ramalina leiodea Nyl. (Nylander, 1888). 

A thorough investigation of specimens of this taxon held at PC and H was undertaken by 
Stevens (1982). It was found that two acid races (viz. boninic acid and cryptochlorophaeic acid) 
were present in taxa bearing this name. Because the material designated lectotype contained 
boninic acid, the species R. boninensis Asah. and R. boninensis f. subcalicariformis were 
reduced to synonomy at that time. 

Krog & Swinscow (1976) commented on the similarity between the East African taxon R. 
maritima and material from Bourbon I. and Mauritius. Specimens from the latter two areas had 
been identified as R. subfraxinea by Nylander (1870). Chemical analysis showed they contained 
boninic acid (Krog & Swinscow, 1976), which was also confirmed in this study (see comments 
under R. subfraxinea var. subfraxinea). Morphologically and chemically R. subfraxinea var. 
leiodea and R. maritima resemble one another but differences in spore size sets them apart, the 
former taxon produces spores in the range 8-16 x 4-5 /xm, whereas R. maritima spores lie in the 
range 18-25 x 5-6 /im. 

Distribution and habitat. Ramalina subfraxinea var. leiodea occurs from latitudes 18°S-29°S 
along the eastern coastline of Australia (Fig. 28), with its most frequent occurrence in the 



208 G. NELL STEVENS 

subtropics. The distribution range overlaps that of R. subfraxinea var. confirmata, a mainly 
tropical taxon which extends into the subtropics so that both taxa are sympatric for a large part of 
their distribution (Fig. 31). 

This taxon occurs on mangroves {Avicennia marina and Rhizophora stylosa) and on coastal 
trees {Callitris columellaris and Casuarina equisetifolia) . 

World distribution. Bonin Is, Marianas Is, Mauritius, Reunion I., AustraUa, Lord Howe I., 
Norfolk I., New Caledonia, New Hebrides, Cook Is (Rarotonga), and Hawaii. 

Queensland: Hinchinbrook I, 18°14'S, 146°50'E, on Ceriops tagal, 1979, GNS (BRIU3479NS); Long I, 
Whitsunday Group, 20°21'S, 148°51'E, on coastal shrub, 1979, A. B. Cribb (BRIU3225NSi); Eimeo, 
21°09'S, 149°09'E, on Ceriops tagal, 1977, GNS (BRIU3155NS); St Lawrence, 22°20'S, 149°32'E, on 
Ceriops tagal, 1976, GNS (BRIU2455NS); Gladstone, 23°51'S, 151°16'E, on Rhizophora stylosa, 1980, 
GNS (BRIU3285NS); Turkey, 24°06'S, 151°38'E, on Rhizophora stylosa, 1975, GNS (BRIU1148NS); 
Burnett Hds, 24°46'S, 152°25'E, on Rhizophora stylosa, 1976, GNS (BRIU4296NS); Toogoom, 25°15'S, 
152°40'E, on Callitris columellaris, 1983, GNS (BRIU4152NS); Urangan, 25°19'S, 152°55'E, on Callitris 
columellaris, 1978, GNS (BRIU2884NS); Kin Kin Ck, 26°13'S, 153° on Casuarina sp., 1979, GNS 
(BRIU3223NS); Noosa Hds, 26°23'S, 153°05'E, on rain-forest tree, 1980, GNS (BRIU3547NS); Moreton 
\, 27°20'S, 153°25'E, on Callitris columellaris, 1980, RWR (BRIU3476NS); Cape Byron, 28°37'S, 
153°36'E, on Acacia sp., 1980, RWR (BRIU3883NS). New South Wales: Yamba, 29°27'S, 153°20'E, on 
Avicennia marina, 1977, GNS (BRIU2122NS); Broomes Hd, 29°37'S, 153°20'E, on Casuarina sp., 1982, 
GA^5(BRIU4311NS). 

var. norsticticaN . Stevens, var. nov. Plate 11, fig. 8. 

Thallus corticolus, viridi-griseus caespitosus rigidus erectus vel subpedulus, 1-5-6-5 cm longus, ramifi- 
catione subdichotoma sparsa vel mediocri; rami lati ad 1-1-5 mm, applanati vel canaliculati, apice 
attenuati; superficies impolita, laevigata vel rugosa et crassa, pseudocyphellis laminalibus linearibus, ad 
fundamenta, tubercularis pseudocyphellis marginalibus, sparsa vel numerosa; soralia nulla. Apothecia 
numerosa, marginalia vel subterminalia, calcarata; discus ad 3-5 mm in diametro, concava vel plana vel 
convexa; sporae ellipsoideae vel raro fusiformae, rectae vel raro curvatae, 9-16(-20) x 3-5-5-5 ^tm. 
Acidum norsticticum ± connorsticticum et usnicum continens. 

Holotype: Australia, Queensland, St Lawrence, on Ceriops tagal, 11 July 1978, G. N. Stevens 2459 (MEL 
1048088!). 

Thallus corticolous, pale green to green-grey, caespitose, rigid erect to subpendulous, 1 ■5-6-5 
cm long; branching subdichotomous, sparse to moderate; branch width 1-1-5 mm, compressed, 
flat or canaliculate, some marginal splitting of the branches between the upper and lower 
surfaces, apices attenuate; surface matt, smooth to rugose and coarse, with or without basal 
pseudocyphellae, tuberculate pseudocyphellae marginal few to many when present; holdfast 
delimited; soraha absent. 

Apothecia common, marginal and subterminal, some spurred; disc 3-5 mm diam., concave, 
plane to convex; margin thin, with or without lateral branchlets; spores ellipsoid rarely fusiform, 
straight, rarely curved, 9-16(-20) x 3-5-5-5 /xm. 

Chemistry. Norstictic acid, ± connorstictic acid, and usnic acid. 

Remarks. This taxon is an Austrahan endemic; examination of esorediate maritime taxa from 
overseas herbaria failed to find any norstictic acid taxon which resembled it. In Austraha the 
taxon closest to it morphologically is R. subfraxinea var. subfraxinea. Both taxa are sympatric 
for part of their distribution and have similar habitats so that the similarity in morphology could 
be due to convergent morphology under the same environmental pressures (Fig. 31). 

Ramalina subfraxinea var. norstictica produces branchlets around the margins of the apothe- 
cia at times, but this is not a constant character and cannot be used taxonomically . Splitting along 
the margins of the branches also occurs, but this is also common in the salazinic acid taxon 
belonging to the R. subfraxinea complex. The rank of variety was given to this norstictic acid 
taxon because of its distinct chemistry and its geographical occurrence along the northern 
Austrahan coastline (Figs 29, 31). 

Distribution and habitat. Ramalina subfraxinea var. norstictica occurs along the northern and 



RAMALINA IN AUSTRALIA 



209 




Fig. 29 Distribution of R. subfraxinea var. norstictica. 



eastern coastlines of Australia (Fig. 29). Its absence from an area between latitudes 17°- ITS on 
the eastern coast is an anomaly since it is found on the northern coastline between these 
latitudes. It is in fact replaced by the salazinic acid taxon R. tropica in this section of the eastern 
coastline. It could be inferred that a change from one depsidone to the other came about with 
change in environmental pressures as norstictic acid is almost certainly a biogenetic precursor of 
salazinic acid (Elix, in litt.). However, no thalli were found containing both norstictic acid and 
salazinic acid. 

The morphology of the norstictic acid taxon is variable and appears to be ecophenotypic, 
ranging from thalli with narrow, canaliculate branches bearing subterminal apothecia, to broad, 
flat, branched thalli bearing large, marginal apothecia. Tuberculate pseudocyphellae occur on 
thalU growing in very dry or polluted areas. The spores of R. subfraxinea var. norstictica are 
intermediate (in both size and shape) between the small ellipsoid spores of R. subfraxinea var. 
confirmata and the large fusiform spores of R. tropica. 

World distribution. Australia. 

Western Australia: Cygnet Bay, 16°35'S, 123°02'E, on Bruguiera sp., 1980, S. Kenneally (PER001045); 
Swan Point, 16°25'S, 123°02'E, 1906, W. V. Fitzgerald (NSW L4117). Northern Territory: Port Darwin, 
12°26'S, 130°48'E, 1890, M. Holtze (MEL 9477); East Alligator R. 12°08'S, 132°40'E, on Ceriops tagal, 
1979, E. Hegerl, (BRIU3114NS); Cobourg Pen., Caiman Ck, 11°25'S, 132°06'E, on mangrove, 1968,7. R. 
Maconochie (MEL100288); Tomkinson R, 12°13'S, 134°16'E, on Avicennia sp., 1975, D. Grace 
(MEL1012777); Melville Bay, 12°15'S, 136°40'E, on Ceriops tagal, 1948, R. L. Specht (SA 97528120); 



210 G. NELL STEVENS 

Groote Eylandt, Hemple Bay, 13°49'S, 136°38'E, on mangroves, 1948, R. L. Specht (SA 97527533); 
Carrington Ck, 15°44'S, 136°36'E, on mangroves, McKay (MEL102350). Queensland: Weipa, 12°38'S, 
141°55'E, on mangroves, 1974, R. L. Specht (BRIU3584NS); Hinchinbrook I, 18°22'S, 146°15'E, on 
Ceriops tagal, 1979, GNS (BRIU3185NS); Townsville, 19°15'S, 146°50'E, on Ceriops tagal, 1975, B. 
Bailment (BRIU3120NS); Bowen, Dry Ck Rd, 20°04'S, 148°22'E, on Ceriops tagal, 1979, GNS 
(BRIU3141NS); Mackay, Eimeo, 21°09'S, 149°09'E, on Ceriops tagal, 1976, GNS (BRIU3434NS); St 
Lawrence, 22°21'S, 149°32'E, on Excoecaria agallocha, 1976, GNS (BRIU2463NS); Corio Bay, 22°58'S, 
150°46'E, on Rhizophora sp., 1975, D. Tarte (BRIU1170NS); Keppel Sands, 23°21'S, 150°47'E, on 
Excoecaria agallocha, 1975, RWR (BRIU760RR); Port Alma, 23°35'S, 150°51'E, on Ceriops tagal, 1975, 
/?IV/?(BRIU1039RR); Gladstone, 23°50'S, \5\° W¥., on Rhizophora stylosa, 1980, GA^5(BRIU3259NS). 



20. Ramalina tropica N. Stevens, sp. nov. Plate 12, figs 1, 2, 6. 

Thallus corticolus et saxicolus, virentes-pallido caespitosus rigidus erectus, usque ad 4-0 cm altus, 
ramificatione subdichotoma sparsa vel mediocri; rami lati ad 0-5-1 -0 mm, subteretes vel teretes vel raro 
planares, apice attenuati curvi; superficies impolita vel nitida, laevigata vel rugosa, pseudocyphellis 
laminalibus linearibus instructa aut sine pseudocyphellis; soralia nulla. Apothecia raro aut vulgariter, 
subterminalia et calcarata vel marginala, ramis flexuosis insidentes; discus ad (1-)1 •5-3-0 mm in 
diametro, planus vel convexus; sporae fusiformae, rectae, ll-18(-22) x 3-8-5(-6) /u,m. Acidum 
salazinicum ± protocetraricum et usnicum continens. 

Holotype: Australia, Queensland, Lizard Island, 26 December 1974, R. L. Specht (MEL 1048087!). 

Thallus corticolous, or saxicolous, pale green, caespitose, rigid, erect, usually to 2 cm high, 
exceptionally to 4 cm; branching subdichotomous, sparse to moderate, branches arising from 
the base with lateral branchlets common; branch width 0-5-1-0 mm, subterete to almost terete, 
rarely flat, never canaliculate, some marginal splitting of the branches between the upper and 
lower surfaces, apices tapering distally, often recurved; surface matt or shiny, smooth to rugose, 
fine linear pseudocyphellae and tuberculate pseudocyphellae sometimes present; holdfast 
delimited; soralia absent. 

Apothecia rare to common, marginal on flexuous branches or subterminal with the branch 
continuing to grow forming a long curved spur up to 10 mm, disc (l-)l-5-3-0 mm diam. , plane to 
convex, some assuming a helmet shape; margin entire; spores fusiform, rarely ellipsoid, 
straight, rarely bent, some appearing as 3-celled; 12-18(-22) x 3-8-5(-6) /am. 

Chemistry. Salazinic acid, ± protocetraric acid, and usnic acid. 

Remarks. The erection of this new species has been made cautiously. There were several 
reasons for making this decision rather than regarding the salazinic acid taxon as another 
chemical variety of the R. subfraxinea complex, (i) Morphologically the four depside taxa and 
the norstictic acid taxon closely resemble one another in producing thalli with flat or canaUculate 
branches, whereas the salazinic acid taxon mainly produces subterete branches and in some 
saxicolous forms, terete branches; (ii) The large size of the spores is in excess of the size 
produced by any of the other taxa in the complex; (iii) the distinct depsidone chemistry. 

Ramalina tropica resembles the East African species R. fecunda Krog & Swinscow, which 
contains salazinic acid and produces large fusiform spores. However, the latter taxon has 
branches which are flat or canaliculate, not subterete (Krog, in litt). 

Other maritime species producing salazinic acid are R. zollingeri Szat. (Java), R. sideriza 
Magnusson (Hawaii), and undescribed material (Fiji). Before any affinities could be established 
between these taxa and R. tropica more material would need to be examined. 



Plate 12 Thallus growth forms in the Australian Ramalinae. Scale in mm. Fig. 1 R. tropica (sterile form). 
Fig. 2 R. tropica (type specimen). Fig. 3 R. subfraxinea var. subfraxinea (top - narrow form; bottom - 
broad form). Fig. 4 R. subfraxinea var. confirmata (broad form). Fig. 5 R. subfraxinea var. norstictica 
(sparse apothecia). Fig. 6 R. tropica (terete, inland form). Fig. 7 R. subfraxinea var. subfraxinea (broad 
form). Fig. 8/?. subfraxinea var. leiodea. Fig. 9R. subfraxineavax . norstictica. Fig. 10/?. subfraxinea var. 
confirmata. 



RAMALINA IN AUSTRALIA 



211 




212 



G. NELL STEVENS 




Fig. 30 Distribution ofR. tropica. 



Distribution and habitat. The distribution of R. tropica is restricted to an area on the eastern 
coasthne of Cape York between latitudes 13°S (Claudie River) and 18°S (Hinchinbrook I.) [Figs 
30, 31]. Because of its restricted distribution range compared with that of the other R. 
subfraxinea taxa, indications are that R. tropica may have certain habitat requirements not 
found elsewhere along the coast. 

Ramalina tropica is both corticolous and saxicolous over its range, occurring both coastally 
and on the coastal hills. Morphologically variable, it produces several morphotypes, e.g. at 
latitude 18°S it is sympatric with the rest of the R. subfraxinea complex and closely resembles 
these taxa, illustrating the phenomenon of convergent morphology. At latitude 16°S it is 
sympatric with R. subfraxinea var. subfraxinea and both produce a straggly, wiry growth form 
with dense lateral branchlets; however, the characteristic rounding of the branches in R. tropica 
is detectable when comparing the thalli with flattened branches of R. subfraxinea var. subfrax- 
inea thalh. 

Corticolous thalli of R. tropica attain their maximum length of 4-0 cm on Lizard Island in the 
mangrove environment where Ceriops tagal is the usual phorophyte. The branches are very 
narrow, subterete, and bear marginal apothecia subtended by markedly attenuate recurved 
branch apices (Plate 12, fig. 1). Ramalina tropica also grows on granite rock on Lizard Island, but 
the saxicolous morph differs from the corticolous morph in producing a rigid, sparsely branched 
thallus, bearing few or no apothecia. 

Inland collections of this taxon were made at Abbey Peak (650 m altitude) at latitude 14°18'S, 



RAMALINA IN AUSTRALIA 



213 




Fig. 31 Distribution ranges of the six acid taxa in the R. subfraxinea complex: var. subfraxinea (sekikaic 

acid race) ; var. subfraxinea (divaricatic acid race) ; var. norstictica ; var. confirmata 

; var. leiodea ; R. tropica — — — . 



where it occurs on windblown shrubs at the summit, and at Lighthouse Mountain (400 m 
altitude) at latitude 16°38'S where it occurs on granite rock at the summit. These morphotypes 
are more terete than coastal thalli and produce a thicker cortex. Increased thickness of the 
cortex is a feature reported by Rundel (1982) as an adaptation to decrease the rate of 
evaporation from the thallus in extreme conditions where selection for low rates of evaporative 
loss is more important than rapid rate of water uptake. The terete branches and fasciculate 
growth form (Plate 12, fig. 6) may be a response to increased moisture stress as discussed by 
Sheard (1978); these adaptations would ensure that less thallus area was effected by heat in an 
environment which is exposed to extreme heat and dryness for long periods. 

World distribution. Australia. 

Queensland: Claudie River, Charlie Taylors Landing, 12°36'S, 143°42'E, on Ceriops tagal, 1983, GNS 
(BRIU4111NS); Abbey Peak, 14°18'S, 144°30'E, on shrub on summit, 1983, C. McCracken 
(BRIU4239NS); Lizard I, 14°41'S, 145°28'E, on granite, 1974, R. L. Specht (BRIU3780NS); Lizard I, 
14°41'S, 145°28'E, on mangroves, 1974, R. L. Specht (MEL1048087) Type; Lighthouse Mt, between Mt 
Carbine & Mt MoUoy, 16°35'S, 145°13'E, on granite, 1983, C. McCracken (BRIU4218NS); Daintree R. 
south branch, 16°18'S, 145°24'E, on Ceriops tagal, 1983, GNS (BRIU4189NS); Mossman Rifle Range, 
16°29'S, 145°25'E, on Ceriops tagal, 1983, GNS (BRIU4188NS); South Mission Beach, 17°56'S, 146°05'E, 
on rain-forest tree, 1983, GNS (BRIU4163NS); Murray R. 18°02'S, 146°03'E, on Ceriops tagal, 1979, D. 
Tarte (BRIU3159NS); Hinchinbrook 1, 18°20'S, 146°15'E, on Ceriops tagal, 1979, GNS (BRIU3160NS). 



214 



G. NELL STEVENS 





— ! 


r- ^^_ 




___ 





J,:^^,,,,,,,rrmmmmummmmmm\\m\mmmm 




Plate 13 Photographs of Austrahan Ramalina types. Scale in mm. Fig. 1 R. australiensis (PC). Fig. 2 R. 
myrioclada (G). Fig. 3 R. exiguella (BRI). Fig. 4 R. glaucescens (M). Fig. 5 R. lacerata (G). Fig. 6 R. 
leiodea war . fastigiatula (G). Fig. 7 7?. calicaris var. australica (H). Fig. 8 /?. &revw (NSW). 



RAMALINA IN AUSTRALIA 



215 





1 , .i^^^^^^iJimliiiiliiiitmi!i(HitfnltiHim,i.n,i„„ii...i..,,|iniHff,(t 



Ctt-rat-Co. \*.tWK W^.t-t.VTa.4 




7:::-:;^7:^-^::::: Iti5^7m=f9 LecTVTYPs: 




Plate 14 Photographs of AustraUan Ramalina types. Scale in mm. Fig. 1 R. farinacea var. squarrosa (G). 
Fig. 2 /?. farinacea var. dendroides (G). Fig. 3 /?. perpusilla (BRI). Fig. 4 R. fissa (G). Fig. 5 R. inflata 
(FH). Fig. 6 R. tasmanica (H). Fig. 7 R. nervulosa var. nervulosa (G). Fig. 8 ^. pacifica (TNS). 



216 




Plate 15 Photographs of Australian Ramalina types. Scale in mm. Fig. 1 R. unilateralis (G). Fig. 2 R. 
subfraxinea (PC). Fig. 3 R. leiodea (PC). Fig. 4 R. celastri (FH-isotype). Fig. 5 R. ovalis (BM). Fig. 6 R. 
fraxinea**R. confirmata (H - isotype). Fig. 7 R. minscula var. alba (WELT). Fig. 8 R. fraxinea**R. 
confirmata (H - lectotype). 



RAMALINA IN AUSTRALIA 217 

Appendix. Disposition of excluded Ramalina taxa 

The position of other Ramalina taxa recorded from AustraHa in Wetmore (1963), and Weber & Wetmore 
(1972) is indicated below. 

R. anceps = R. australiensis Nyl. or R. filicaulis N. Stevens 

R. angulosa = R. australiensis Nyl. 

R. calicaris = R. glaucescens Krempelh. 

R. calicaris var. canaliculata = R. glaucescens Krempelh. 

R. complanata = R. subfraxinea Nyl. 

R. confirmata = R. subfraxinea var. confirmata (Nyl.) N. Stevens. 

R. dendriscoides = R. tenella Miill. Arg. or R. peruviana Ach. 

R. dendriscoides var. minor = R. tenella Miill. Arg. or R. peruviana Ach. 

R. dilacerata var. alba = R. inflata subsp. perpusilla (Stirton) N. Stevens 

R. ecklonii = R. celastri (Sprengel) Krog & Swinscow 

R. ecklonii var. membranacea = R. celastri (Sprengel) Krog & Swinscow 

R. ecklonii var. ovalis = R. celastri subsp. ovalis (J. D. Hook & Taylor) N. Stevens 

R. ecklonii var. tenuissima = R. celastri (Sprengel) Krog & Swinscow 

R. fastigiata = R. glaucescens Krempelh. 

R. fraxinea = R. glaucescens Krempelh. 

R. fraxinea var. ampliata = R. glaucescens Krempelh. 

R. fraxinea f. platyna = R. fissa (Miill. Arg.) Vainio 

R. fraxinea var. taeniata = R. glaucescens Krempelh. 

R. furcellata = R. australiensis Nyl. 

R. geniculata = R. inflata J. D. Hook. & Taylor 

R. geniculata var. olivacea = R. inflata subsp. perpusilla (Stirton) N. Stevens 

R. gracilenta = R. exiguella Stirton 

R. gracilis = R. exiguella Stirton or R. filicaulis N. Stevens 

R. homalea = unknown? 

R. inflata var. gracilis = R. inflata (J. D. Hook. & Taylor) J. D. Hook. & Taylor 

R. intermedia = R. inflata subsp. australis N. Stevens 

R. javanica = R. peruviana Ach. 

R. linearis = R. celastri (Sprengel) Krog & Swinscow 

R. pollinaria = R. unilateralis F. Wilson 

R. polymorpha = unknown 

R. polymorpha f . emplecta = unknown 

R. pusilla = R. inflata subsp. australis N. Stevens 

R. scopulorum = R. subfraxinea Nyl. var. ? 

R. scopulorum var. cuspidata = R. subfraxinea Nyl. var. ? 

R. scopulorum var. subfarinacea = R. pacifica Asah. 

R. tayloriana = R. nervulosa var. luciae (Molho et al.) N. Stevens 

R. usnea = R. australiensis Nyl. 

R. usneoides = R. australiensis Nyl. 

R. yemensis = R. celastri (Sprengel) Krog & Swinscow 

R. yemensis var. ovalis = R. celastri subsp. ovalis (J. D. Hook. & Taylor) N. Stevens 

Acknowledgements 

This research was carried out for a Doctor of Philosophy thesis at the University of Queensland Botany 
Department. Some funding was provided from University Research Grant Funds. I thank my supervisors 
Dr R. W. Rogers and Professor H. T. Clifford for their discussions and guidance during the course of this 
research, and Dr J. A. Elix for his generous help with chemical problems and Mr R. B. Filson for his help 
and encouragement. I wish to thank all overseas lichenologists with whom I corresponded during this 
study, especially Professors H. Krog and D. L. Hawksworth, Drs M. E. Hale and D.J. Galloway, and Mr 
P. W. James. 

I would also like to thank the directors and curators of the following herbaria for supplying material on 
loan: AD, BM, BRI, CBG, EDIN, FH, G, HO, LD, M, MICH, MEL, NSW, O, PC, PER, S, TNS, TUR, 
UPS, UWA, US, and W. In addition I am grateful to the following persons who placed their private 
collections at my disposal: C. Smith (Hawaiian RamaHnae), G. & B. Hayward (New Zealand Ramalinae), 
H. Osorio, M. Pinto (South American Ramalinae), J. Clarkson (Torres Strait Ramalinae), R. Seppelt 



218 G. NELL STEVENS / 

(Macquarie Island Ratnalinae), R. Specht, J. Berjak (South African Ramalinae), G. Kantvilas (Tas- 
manian Ramalinae) and A. Archer, E. Dahl, J. Elix, R. Rogers, N. Sammy, C. Scarlett, H. Streimann 
(mainland Australia Ramalinae). I am grateful to Dr A. Henssen for supplying two of the photographs 
used in this paper. All SEM and TEM photographs were taken by Mr J. Hardy of the Electron Microscope 
Centre, University of Queensland. I thank M. Lamy for supplying information from the Verreaux 
journals. 

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RAMALINA IN AUSTRALIA 219 

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222 



G. NELL STEVENS 



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Index 



Accepted names are in roman and synonyms in italic; new names and principal references are in bold. 
An asterisk (*) denotes a figure. 



Alectoria 116, 118, 119, 147 

anceps 147, 160, 217 

angulosa 217 

arabum 116 

attenuata 158 

australiensis 119, 124, 127, 129, 

138,143,149,150,151,152, 

153M54M55,214*,217 
Bistortae 1 16 
Bitectae 116 
boninensis 207 

f . subcalicariformis 207 
bourgaeana 129 
brevis 120, 163, 166, 214* 

var. brevissima 120, 193, 194 
Buibothrix 146 
caespitella 121, 123*, 129, 138, 139, 

143, 149, 150, 153*, 155, 156* 
calcarata 146, 186, 194 
calicaris217 
f . Ecklonii IQH 

var. australica 120, 163, 166, 214* 
var. canaliculata 217 
var. ovalis 184 
canariensis 121, 123*, 124, 129, 

132, 138, 139, 143, 149, 150, 

151, 153*, 156, 157*, 178 
capitatal21,123*,130, 179 
carpathica 115 
celastri (Parmelia) 180 
celastri (Ramalina) complex 124, 

129 
subsp. celastri 128*, 129, 131*, 

136, 138, 139, 143, 149, 150, 

151,171,180, 181*, 182, 183*, 

184, 216*, 217 
subsp. ovalis 108, 129, 131*, 138, 

139,143,149,150,152,181*, 

183*, 184, 217 
Cenozosia 115, 116, 117, 118, 119 
ceruchis 115, 116 
chondrina 160, 161 
Ciliatell6 
combeiodes 115 
complanata 115, 126, 217 
Compressiusculae 115, 116 
confirmata 119 
consanguinea 146, 189 
continentalis 182 
Corticatae 116 
cumanensis 182 
curnowii 130 
decipiens 129 



dendriscoides 176, 217 

var. minor 217 
Desmazieria 115, 116, 117, 119 
Dieverniain, 119, 129 
dilacerata var. alba 217 
Dirina 147 
Dirinaria 148 
dumeticola 197 
duriaei 129 

ecklonii 180, 182, 184, 207, 217 
var. lobulifera 182 
var. membranaceae 182, 217 
var. ovalis 184, 217 
var. tenuissima 182, 217 
Ecorticatae 116 
Ellipsosporae 116 
Euramalina 115, 116 
evernioides 116 

exiguella 119, 120, 128*, 129, 131, 
136,138,143,149,151,153*, 
154, 158, 159*, 177, 214*, 217 
farinacea 115, 124, 126, 129, 130*, 
132, 134, 147, 197, 198, 200, 
201 
var. dendroides 120, 170, 172, 

215* 
var. nervulosa 120, 195, 197, 200 
var. squarrosa 120, 170, 172, 
215* 
fastigiatall5,217 
fastigiatus (Lichen) 117 
fecunda 210 

filicaulis 108, 119, 124, 129, 130*, 
136,138,143,149,151,153*, 
154,160,161,217 
fimbriata 119, 121, 122*, 123*, 129, 
132,138,139,140,141,143, 
149, 150, 152, 153*, 161, 162*, 
178 
fissa 131, 132, 138, 143, 149, 150, 
151,152, 157, 181*, 185, 191, 
193, 194, 195*, 215* 
Fistularia Vainio 115, 116, 126 
Fistulariae R. Howe 116 
Fistulariella Bowler & Rundel 110, 

115,116,117,119 
Fistulosae Hue 115 
flaccescens 115 
fraxinea 115, 150,203,217 
p yemensis 180, 182 
f. platynal94,217 
subsp. confirmata 205 
var. ampliata 217 



var. brevissima 193 
var. taeniata 217 
furcellata 217 
Fusisporae 116 

geniculata 117, 185, 186, 194, 217 
var. compacta 120, 188 
var. olivacea 217 
glaucescens 119, 129, 131, 133, 138, 

139, 141*, 143, 149, 152, 163, 

164*,165, 166*,167, 168, 

214*, 217 
gracilentia 217 
gracilis 119, 158,217 
homalea 115, 116,217 
implectens 147 
inanis 115, 116 
inflata (Cetraria) 185, 215* 
inflata (Fistulariella) 117, 119, 185, 

187, 194 
inflata (Ramalina) complex 129, 

130*,133, 139, 191,192 
subsp. australis 108, 138, 139, 

143,149,150,151,152,181*, 

186*, 191, 193, 194,217 
subsp. inflata 128*, 130, 131, 138, 

139, 143, 146, 149, 150, 152, 

181*,183, 185, 186*,187,217 
subsp. perpusilla 108, 128*, 130*, 

136, 138, 143, 146, 149, 150, 

152, 159, 181*, 183, 186*, 187, 

188, 190, 217 
var./i5jal20, 193 
var. gracilis 217 
insularum 201 
intermedia 217 
intermediella 172, 200, 201 
javanica 217 
knightiana 120, 185 
lacerata 120, 163, 214* 
laevigata 182 

leiodea 117, 148, 203, 207, 216* 
\ar . fastigiatula 119, 163, 165, 

167, 214* 
leptocarpha 147 
linearis 115, 182, 271 
litorea 128*, 129, 136, 138, 143, 

149, 151, 153*, 169, 170* 
luciae 199 
maritima 207 
mediterranea 191 
melanothrix 115, 158 
microspora 169 
minuscula var. alba 120, 188, 216* 



RAMALINA IN AUSTRALIA 



223 



Myelopoea 115, 116 

Myelopoeae 116 

myriodada 120, 152, 154, 214* 

nervulosa var. dumeticola 121, 
122*, 123*, 124, 134, 136, 138, 
143,149,151,196,197,198, 
199*, 201 
var. Iuciael21,124, 125, 134, 
136,138,143,149,151,177, 
196, 199*, 200, 201, 217 
var. nervulosa 121, 124, 125*, 
129,136,138,143, 149,151, 
177, 195, 196, 198*, 200, 215* 

Nieblall5,116, 117, 119, 129 

ovalis 184, 216* 

pacifica 121, 124, 125*, 127*, 129, 
132,136,138,143,149,151, 
159,177,196,200,201,202*, 
215*, 217 

Parmelia sect. Hypotrachyna 147 

Parmelina 146 

perpusilla 120, 188, 215* 

pertenuis 158 

peruviana 115, 120, 121, 122*, 124, 
125,129,131,136,138,143, 
149,151,154,170,171,172, 
173*,174, 176, 198,200,217 

Physcia 147 

pollinaria 115, 161, 178,217 

polymorpha 115, 217 
f. emplecta 217 

populina 130 

Pseudocyphellaria 146 

Pseudoparmelia 146 

Psoroma 146 



pumila 189 
pusillal86, 191,217 
Ramalinal09*,115, 149 
Ramalinaceae 115, 116, 117, 118, 

119 
Ramalinopsis 117, 119 
reducta 121, 122*, 123*, 129, 132, 
138, 139, 141*, 143, 149, 151, 
171, 175*, 176, 183 
Relicina 146 
rigida 158 

scopulorum 115, 217 
var. cuspidata 217 
var. subfarinacea 217 
sideriza 210 
siliquosa 133, 134 
sinaloensis 126 
Solidae 1 16 

Speerschneidera 117, 119 
sprengelii 182 
Sterecaulon 146 
strepsilis 130, 179 
subfraxinea 203, 204, 207, 216 
complex 129, 134, 146, 148, 208, 

210,217 
subsp. leiodea 207 
var. confirmata 108, 125*, 127*, 
136, 138, 143, 149, 152, 159, 
176, 196, 202, 205, 206*, 208, 
211, 213, 216*, 217 
var. leiodea 108, 127*, 136, 138, 
143,149,152,159,196,203, 
205, 206*, 207, 211, 213 
var. norstictica 108, 127*, 136, 
138,143,149,151,196,208, 



209*, 211, 213 
var. subfraxinea 125*, 127*, 136, 

138, 143, 148, 149, 152, 196, 

203, 204*, 207, 208, 211, 212, 

213, 216*, 217 
subgeniculata Knight 120, 185 
subleptocarpha 147 
subpusilla 194 
tasmanica 117, 185, 215* 
tayloriana217 
tenellal21,122*,123*,129, 132, 

136,138,143,149,151,171, 

172,176, 177*, 217 
Tenuicarticatae 116 
Tenuicorticate 116 
Teretiusculae 115, 116 
testudinaria 115 
Trichoramalina 115, 117, 119 
tropica 108, 136, 138, 143, 149, 150, 

151, 209, 210, 211, 212*, 213 
Tubulosae 116 
unilateralis 120, 121, 122*, 123*, 

124, 129, 132, 138, 139, 141*, 

143,149,151,156,157,161, 

171,177, 178*, 216*, 217 
usnea 108, 147, 154, 160, 217 
Usneall6, 118, 119, 146 
Usneaceaein, 118, 119 
usneoides 154, 217 
whinrayi 130*, 132, 133, 138, 139, 

143,149,151,171,179,180* 
yemensis 180, 182, 184, 217 

var. ovalis 184, 217 
zollingeri 210 



British Museum (Natural History) 

Ferns of Jamaica 

A guide to the Pteridophytes 

G. R. Proctor 

This flora records and describes the 579 species and 30 varieties of ferns occurring in 
Jamaica. The succinct species descriptions include relevant synonymy and incorpor- 
ate distributional data both within and outside Jamaica. Special emphasis is given to 
the subtle distinctions between closely related species and all genera are illustrated. 
Keys to the genera and species facilitate a wider use of the flora in the West Indies and 
northern South America. The author, one time Senior Botanist in charge of the 
Herbarium of the Science Museum, Kingston, Jamaica, is an outstanding field 
botanist and his expertise is reflected in the practicality of the flora and especially 
in the habitat and ecological information. This volume represents an important 
addition to our knowledge of the flora of the West Indies. 

1985, 631pp, 135 hne illustrations, 22 maps. Hardback. 
565 00895 1 £50.00 



Titles to be published in Volume 16 



studies in the genus Hypericum L. (Guttiferae) 
7. Section 29. Brathys (part 1) 
ByN. K. B.Robson 

The lichen genus Ramalina in Australia 

By G. N. Stevens 

An annotated list of vascular plants collected in the valleys south of 
Mt Everest 

By G. Miehe 

Title to be announced 



Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk 
Printed in Great Britain by Henry Ling Ltd, Dorchester 



B'"-^' 



Bulletin of the 

British Museum (Natural History) 



An annotated list of vascular plants 
collected in the valleys south of 
Mt Everest 

Georg Miehe 



Botany series Vol 16 No 3 27 August 1987 



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

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WorWLw^ abbreviation: Bull. Br. Mas. nat. Hist. (Bot.) 



© British Museum (Natural History), 1987 



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

Editor of Bulletin: Mr J. R. Laundon 

Assistant Editor: Dr A. J. Harrington 

Editor's Assistant: Miss M. J. Short 



ISBN 565 08016 4 
ISSN 0068-2292 



British Museum (Natural History 
Cromwell Road 
London SW7 5BD 




Botany series 

Vol 16 No 3 pp 225-268 



Issued 27 August 1987 



, (M*T' 

An annotated list of vascular plants collected in the 
valleys south of Mt Everest 

Georg Miehe^ 

Department of Geography, University of Gottingen, Goldschmidtstrasse 5, D-3400 
Gottingen, West Germany 

Contents 

Synopsis 225 

Introduction 225 

Deliminationofthearea 225 

Outline of the climate 226 

Altitudinal belts of vegetation 228 

Remarks on timber-line ecology 231 

Remarks on the influence of grazing, burning, 

and wood-cutting 232 

Alphabetical list of vascular plants, their life forms, 

associated formations, and records of collection 233 

Key to plant life form symbols 233 

Key to plant formation symbols 234 

Abbreviation list of collectors' names 235 

Alphabetical list of flowering plants 235 

Alphabetical list of ferns and fern allies 261 

Acknowledgements 263 

References and other relevant literature 263 

Synopsis 

An alphabetical checklist has been compiled, based mainly on collections in the British Museum (Natural 
History) , of plants at altitudes above 2300 m in east central Nepal, in the upper catchment of the Dudh Kosi 
and the transverse valleys to the south of Mt Everest, eastwards to 87° lO'E. At least 1020 species of 
phanerogams have been collected in this area. Information is provided on life forms and associated plant 
formations, and voucher specimens are cited to vaUdate the records. Introductory notes are provided on 
the altitudinal zonation of the vegetation and on the tree-line. The cloud forests on the humid southern side 
of the main Himalayan range have a potential (natural) cUmatic tree-line which is at c. 4400 m on sunny 
slopes and 200 m lower on shady slopes, as is typical for humid mountains north of the tropics. The upper 
catchment of the Dudh Kosi, which is known as the Khumbu Himal, has been deforested on the sunny 
slopes by the Sherpas, who colonized these valleys from above, by traversing the main range 400 years ago 
and settling permanently, thereby causing widespread secondary vegetation due to extensive uncontrolled 
grazing. 

Introduction 

Delimitation of the area 

The valleys south of Mt Everest, Nepal, included in this account of the flora, are shown in Fig. 1 . 
The only ecologically marked delimitation is the northern boundary which is formed by the main 
Himalayan range. On the northern side there is an arid Tibetan flora, while the south is taken to 
be a part of the humid east Himalayan flora. The western boundary is the watershed between the 
Rolwaling valley and the south-west ridge of Mt Numbur, i.e. the ridge to the west of the 
Lumding Khola. The north-eastern boundary is less definite, nearly coinciding with the eastern 
extension of the Khumbu Himal map of Erwin Schneider (1965). It includes the alpine belt of the 
Barun Khola, with all records which were clearly above the upper cloud forest. The altitudinal 
limit is c. 3800 m at about 87° lO'E, south of Chamlang, Amai Dablang, and Kang Taiga; the 



Bull. Br. Mus.nat. Hist. (Bot.) 16 {3): 225-268 Issued 27 August 1987 



226 



G. MIEHE 



• miin settltmtnt 
o summer settlemtnt 

Cob indptssis 

Glacier 




Fig. 1 The valleys south of Mt. Everest. 



Hinku Drangka (Inugkhu), Hunku Drangka (Hongu), and Iswa Khola run to the south, to join 
the Arun valley in the east or the Dudh Kosi in the west. Only a few botanists have visited these 
valleys and their records all belong to the alpine belt and the upper cloud forests. 

The vast majority of records are from along the route to Mt Everest. In this area, where the 
Dudh Kosi valley leads down from the main range between Cho Oyu in the west and Lhotse in 
the east, the southern . limit of the records assembled here is drawn at 2300 m, south of 
Chaunrikharka. The Dudh Kosi gorge south of Namche Bazar is known as Pharak, while the 
higher catchment of the Dudh Kosi north of 27° 27'N is called Khumbu. The area covered is 
rather heterogeneous in its floristic pattern. The south-eastern valleys, the Barun Khola and the 
Iswa Khola, are likely to prove much more humid (like the east Himalayas) than the Khumbu 
Himal. The latter was classified by Stainton (1972) as belonging to the 'inner valleys', as it shows 
a degree of rain-shadow effect. The other component contributing to the heterogeneity is 
man-made, because the valleys of the Khumbu Himal have been cleared to a much greater 
extent than is typical for the upper cloud forest belt of the southern slope of the main Himalayan 
range. The more arid and continental aspect may be due in part to the presence of ruderals and 
weeds, thus reflecting the influence of man. 

Outline of the climate 

Climatic data in mountain areas are usually inadequate. All meteorological stations are typically 
situated near settlements, and these are mostly located on valley floors or on less cloudy sites. 
Thus, for most mountain areas, the data available give the minimum possible rainfall, and are 
valid only for that small strip on the valley bottom, covering less than 5% of the area. 

The data available are summarized in the climatic diagram (Figs 2-4) after Gaussen as drawn 
by Walter & Lieth (1966), which takes the temperature line as an approach to evapo- 
transpiration. The relation between temperature and rainfall is simplified as 10°C = 20 mm, 
taken from the mean temperature and the mean monthly rainfall, which is reduced to one-tenth 



Chaunrikharka (2743m) 

(mean monthly temperatures 
of Namche Bazar +0,58^/ 
IOOfn/1949-1962 



PLANTS SOUTH OF MT EVEREST 
2283,6mm 



227 



c Namche Bazar {3440m) ca. 6,6° 1048 mm 




Fig. 2 



Month* Source^ Dhw 6 Nmyimii 1965. Josh 1981 



'Lhajung' (4420m) 
(27°53,6' N/86°49'7" E.Gr.) 

(1974,1974) 



I II III IV V VI VII VIII IX X XI XII 

Months Souice: N Dhat. J Naiayarun 1965. 



Fig. 3 



-1.27"C 
528mm 




SoiHce K Higudii lEdl 1984. Aw 3 



Fig. 4 
Figs 2-4 Climatic diagrams . After Walter & Lieth (1966) . 



for graphical reasons, if more than 100 mm. Where the temperature Une lies above the rainfall 
Une it is relatively arid, whereas in the opposite case it is relatively humid. 

From the available data (Dhar & Narayanan, 1965) it is probable that the precipitation 
decreases from the periphery of the mountain area to the centre, roughly by 50% for every 12 
km, but this decrease is also combined with a change in altitude. Thus the decrease is nearly 
uniform every 1000 m between 2400 and 4400 m. Since the temperature is decreasing also, it 
does not become drier. 



228 G. MIEHE 

Only in the medium cloud forest belt, at Chaunrikharka is there a dry season in winter, 
although the precipitation is more than double that at Namche Bazar. At Chaunrikharka 86% of 
the precipitation is concentrated in the monsoon period, whereas Namche Bazar receives only 
74% summer rain. At Namche Bazar, situated in the upper cloud forest belt at 3440 m, relatively 
dry months may still occur independently of the overall annual rainfall (1961: 921 mm, with no 
dry month; 1953: 1710 mm, with March and April dry). The number of rainy days per month is 
higher in Namche Bazar, except in June and July, but fewer millimetres per day fall than in 
Chaunrikharka. The distribution of heavy precipitation, especially snowfall, is of some econ- 
omic importance. At Namche Bazar, the months of heavy 24 hour to 72 hour precipitation are 
those of sowing and harvesting, thus especially before the introduction of the potato (Fiirer- 
Haimendorf, 1964: 8-11), the harvest, which was then of buckwheat and barley, was always 
uncertain. 

Climatic information is also provided by the vegetation, even though we have no exact 
calibration. Closed plant communities, the presence of epiphytes (especially obligate species 
that only grow on trees or rocks), and a dense lichen cover can be interpreted as an indicator for 
humid high-mountain conditions. 

Temperature data have rarely been published. For Namche Bazar (3440 m), the mean 
temperature of the coldest month (January) is -0'4°C, while the mean temperature of the 
hottest month (July) is 12°C (Joshi, 1982: 400). In the alpine belt at 4420 m, the mean 
temperature in January is -7-7°C and that of July is nearly 10°C, over a four year period 
(Higuchi, 1984: appendix). 

Valley winds blow every day, setting in mostly between 10 and 11 a.m. and dropping before 
midnight. They are not as strong as in the transverse gorges like the KarnaU or the Kali Gandaki 
in west and west-central Nepal, but both areas show the same type of clouds, which cover the 
upper slopes of both sides of the valleys with strips, while the sky over the valley bottoms 
remains blue. 

In monsoon and late monsoon conditions, a closed cloud cover from the south often sweeps 
into the valleys, nearly as far as closed forests occur, but above 4200 m only strips of cloud on 
each side of the main valleys advance further up. The valley winds obviously effect the 
wind-exposed vegetation strongly, as indicated by treeless areas and wind-sculptured trees. 

Altitudinal belts of vegetation 

The valleys south of Mt Everest are mostly of the same type as those of the European Alps, being 
typical of high mountains with adequate rainfall to the north of the tropics. Therefore altitudinal 
limits should be expected to be governed by temperature, decreasing at the rate of 5-8°C per 
1000 m and not caused by seasonal drought. This humid type of high mountain is indicated by the 
interlacing of the belt of glaciers down into the free solifluction belt and down into the alpine 
belt. The most striking demonstration of that classification is that the free solifluction belt is 
squeezed into a narrow zone between the glaciers and the vegetation cover of the alpine belt. 
Thus the upper Umit of the characteristic semi-arid altitudinal zone of the high mountains is 
covered with ice, while the potential lower limit of the solifluction process is suppressed by the 
closed vegetation cover of forests and alpine scrub (Kuhle, 1978). All belts are interlocked both 
through natural factors and man's activities. The forest belt is widely cleared and replaced by 
predominantly alpine associations, and 200 m above the present upper forest line, isolated trees 
are found growing in alpine scrub. 

The alpine belt can be roughly divided into a lower zone of moist alpine scrub (dwarf 
rhododendrons and prostrate junipers) and an upper zone with Kobresia pygmaea mats. 
Nevertheless, in wind-exposed situations, the Cyperaceae dominated pastures extend down to 
3000 m, which is at the maximum about 800 m below the potential upper tree-line, and possibly 
the result of initial clearance by fires. Dwarf Rhododendron nivale cushions are found near the 
upper limit of the alpine belt at 5500 m. Hence the altitudinal belts are only differentiated by the 
dominance and not the exclusiveness of their characteristic features. 

The lowest altitudinal belt is the forest belt which extends up to roughly 4200 m. It is 
characterized by epiphytes and should, therefore, be classified as cloud forest. There are both 



PLANTS SOUTH OF MT EVEREST 229 

obligate and facultative epiphytes and the latter can be regarded as indicators of a humid 
climate: creeping ericaceous shrubs are confined to moist soil and are only found on moss- 
covered tree trunks when this substratum is almost constantly damp through rain and fog. Some 
of the cloud forest epiphytes even invade the alpine belt, growing in crevices surrounded by 
cushion-forming alpines, and certain species, mostly found around flushes {Kobresia 
schoenoides, Rhodiola spp.), sometimes even become chasmophytic. 

Mosses are the dominant epiphytes of the forest belt up to 3000 m. They cover trunks and 
branches and are colonized by small phanerophytes as facultative epiphytes. Epiphytic ferns and 
beard lichens occur, but are not dominant. The minimum rainfall in this forest belt is unlikely to 
be less than 1000 mm per year. This medium cloud forest belt is roughly identical with the Tsuga 
dumosa forest and the upper temperate mixed broad-leaved forest described by Stainton (1972: 
1-11,98-100). 

Above 3000 m the upper cloud forest of these transverse valleys is somewhat differentiated 
with regard to dominant tree species depending on the exposure. This is possibly caused by 
decreasing rainfall, which is here probably less than 1000 mm. Forest of Abies spectabilis still 
occurs on south-facing slopes (more those to the south-west than the south-east) and on wind 
exposed ridges on northerly slopes, whereas Betula utilis, which quite often forms an understory 
in Abies forests, becomes dominant on northerly slopes. Rhododendron thickets form the shrub 
layer in fir and birch forests and also, above the upper limit oi Abies and Betula, dominate to 
form pure evergreen thickets ranging from four metres down to one metre near the timber-line, 
representing a transition zone from forest to alpine scrub. 

In this upper cloud forest belt, branches may still be covered with mosses and ferns, but beard 
Uchens are more frequent than in the medium cloud forest belt. On shady slopes, Usnea 
longissima is apparently more widespread and longer than on sunny slopes; branches growing 
out into the valleys are more densely covered with beard lichens. 

Whereas birch forests and the remnants of fir forests are concentrated on the shady and sunny 
slopes respectively. Rhododendron thickets are apparently limited to those slopes which are well 
protected from the wind, but are not "confined to any particular exposure to the sun. 

The alpine belt extends from about 3400 m (owing to forest clearance) up to a maximum 
altitude of 5500 m and is characterized by an almost closed layer of dark alpine soil. 

The most widespread plant associations between 4100 and 5500 m are dwarf shrub communi- 
ties of Rhododendron setosum, R. anthopogon, and R. nivale, with mats of Kobresia pygmaea. 
Wind-blown and south-facing slopes up to 5000 m are dotted with dwarf prostrate junipers. The 
lower Umit of the alpine belt is relatively clear cut on the shady slopes protected from the wind, 
where Rhododendron campylocarpum and R. fulgens mark the upper limit of the forest belt at 
about 4200 m. The sunny slopes are mostly cleared, and between 3400 and 4400 m forest and 
alpine species are intermixed to form the winter grazing grounds. Thus there is a transition zone 
of nearly 1000 m which could possibly be divided into village grazing ground and alpine pastures 
by the presence or absence of prostrate Cotoneaster spp. which are abundant around the 
villages. 

The potential temperature limitation of the alpine belt is thought to be at about 5500 m. For 
the most part this is limited by other factors such as late ice melting or constant falling of frost 
debris from upper slopes. In the upper part of the alpine belt, the cliffs and boulders of white 
granite which have been free from glaciers for long enough are covered with a black carpet of 
Uchens. Clearly the growth of lichens, even though fairly slow, occurs at a faster rate than the 
creation of fresh rock faces by frost action. It can therefore be assumed that with these rocks at 
this altitude, the process of frost cracking is negligible, a controversial conclusion because this 
belt has always been characterized by the dominance of periglacial processes. 

In this upper alpine belt, three herbaceous communities can be distinguished on the alpine 
soil. The first occurs roughly between 4600 and 5000 m on sunny slopes exposed to the wind. 
Kobresia pygmaea provides about 10-30% of the plant cover, but more than 30% is covered by 
flat cushion-forming species such as Anaphalis cavei, Leontopodium monocephalum, L. 
brachyactis, and Arenaria bryophylla. In the second community, which occurs above 5000 m, 
Kobresia pygmaea covers only 5% and the area is dominated by hemispherical cushions such as 



230 G. MIEHE 

Arenaria polytrichoides , by lichens covering the dead remains of plants (one third of which are 
usually dead), and the dark humid soil. The third community consists of nearly 100% cover of 
living Kobresia pygmaea mats, and is confined to depressions between the valley slopes and the 
lateral moraines of the glaciers. These depressions are beheved to be younger than the other 
biotopes on the evidence of the depth of the alpine soil and the diameter of the lichen 
Rhizocarpon geographicum. The problem for all these communities is that the alpine soil was 
obviously established by Kobresia pygmaea, but today this member of the Cyperaceae has 
nearly disappeared except in the depressions along the lateral moraines. Further research is 
needed to decide whether this development has coincided with a climatic change to drier or 
colder conditions, or if it is simply a feature of the succession that at a certain age of alpine soil 
cushion plants become dominant. 

The lower alpine belt is typically covered by moist alpine scrub (as described by Stainton, 
1972: 128). Cushions of Rhododendron anthopogon, R. setosum, R. nivale, and junipers with 
prostrate growth are characteristic of this zone and possibly represent the climax vegetation, 
whereas Lonicera spp. , Spiraea arcuata, Hippophae tibetana, Potentilla fruticosa var rigida, and 
shrubby willows are confined to moist ravines or are part of a plant succession on moraines. 

The dwarf prostrate junipers seem to prefer sunny, wind-exposed slopes, but are more 
frequent on the shady lee slopes when Rhododendron setosum and R. anthopogon are absent. It 
thus seems quite reasonable to suggest that the junipers are restricted by competition to the 
sunny, wind-exposed slopes. 

The cushion-forming rhododendrons show a change in biotope as is explained by the 
ecological law of 'Relative Habitat Constancy and Changing Biotope' (Walter & Walter, 1953). 
At 3480 m and probably 900 mm annual precipitation, R. setosum occurs on sunny, wind- 
exposed slopes; at 4400 m its evergreen cushions are confined to wind-blown but flat sites, 
whereas sunny wind-exposed slopes are free from R. setosum and only R. nivale covers the steep 
parts of the solifluction terraces, while R. setosum is restricted to the shady lee slopes where the 
precipitation amounts to about 500 mm per year. Obviously these rhododendrons change 
biotopes into the ecological niche of a wind-sheltered slope, where the extra-zonal advantage 
compensates for the zonal disadvantages. From eco-physiological experiments carried out by 
Larcher (1963) with R. ferrugineum, it is fairly certain that the Himalayan dwarf evergreen 
rhododendrons are highly sensitive to frost-drought, exacerbated by wind. Thus the daily valley 
winds provide the limiting factor for the distribution of dwarf rhododendrons in the alpine belt, 
but only do so in association with temperatures below 0°C and in the absence of snow protection. 
During the moist summer months the rhododendrons are quite exposed to winds and form 
wind-deformed cushions but in winter they are protected in low ravines and other wind sheltered 
depressions, hence, in the alpine belt of the Khumbu Himal the distribution of dwarf rho- 
dodendron cushions, prostrate junipers, and mats of Kobresia pygmaea depends on exposure to 
the wind, as determined by micro-rehef. Thus vegetation patterns in the alpine belt of these 
subtropical mountains at 28°N are more similar to those found in the Alps at 47°N than would be 
expected. 

The free solifluction belt above the limit of plant cover on alpine soil extends to the maximum 
altitude of thawing (Kuhle, 1978) which is at about 7100 m, according to a record of a Lecidea sp. 
on the south wall of Mt Makalu (4th Jugoslavian Makalu Climbing expedition, pers. comm. J. 
Poelt) which must have been dependent on water. The highest records of flowering plants 
(species unknown) are from c. 6300 m on Cho Oyu (H. Heuberger, pers. comm.) and at 6350 m 
in the western Cwm between camps 4 and 5 by Zimmermann and Hofstetter (Zimmermann, 
1954-1956) on the south side of Mt Everest. Both records refer to small cushions in crevices. 
This biotope provides conditions that are not typical for these altitudes, because daily movement 
of the substratum by thawing and freezing during the vegetative period is the most decisive 
limiting factor for plant colonization, but these processes are minimal in crevices densely stuffed 
with cushions. At the highest altitudes that I was able to find plants (5690 m) it was Saussurea 
simpsoniana that was growing in dense cushions in these crevices. Here competition is often very 
strong, in contrast to the surrounding free solifluction areas which are almost devoid of plants. 
The rather excessive movement of the substratum is only tolerated by a few specialized plants. 



PLANTS SOUTH OF MT EVEREST 231 

which have shoots that lengthen with the downward creeping debris. Probably the shoot length 
of one growing season is approximately equivalent to the distance covered by the debris. Plants 
adapted to this unstable ground might be called 'solifluction acrobats' and include Gentiana 
urnula, Eriophytum wallichii, and Veronica lanuginosa. All other plants are confined to small 
flat patches protected from the stronger debris movement. There may occur here a nearly closed 
cover of plants, small cushions of crucifers, loose flat cushions of Stellaria decumbens, mostly 
with one half dead, and grasses with huge, dense root systems. Typically, their innovation buds 
are not at the surface, as would be expected for hemicryptophytes, but have sunk into the debris 
which has already moved on after the end of the growing period and are thus well protected 
during winter. The flora of the free solifluction belt is quite distinct from the one that occurs in 
patches of debris, ravines, or on gravel along streams in the alpine belt, and only a small number 
of species occur on both sides of the main Himalayan range. 

Remarks on timber-line ecology 

The striking feature of forest distribution in the valleys to the south of Mt Everest, mainly in the 
Khumbu Himal, is that forests predominate on the shady slopes, while the sunny slopes are 
covered with open woodland or grazed open pastures. In the same way, the observation that on 
flat ground, or on one side of ravines, forests grow higher up than on the other slope, suggests 
that open sunny slopes may not be favourable for forests and that this might be the result of 
drought. Yet one can always find solitary trees or groves on the sunny slopes. If these trees or 
groves do not depend on special tectonic conditions such as fault springs, which can be ruled out 
if the herb layer does not indicate moist conditions, one would expect woodlands or forests to 
occur where the topoclimatic conditions are tolerable for a single tree. The conclusion must 
therefore be that single trees or groves on sunny slopes are forest remnants that have survived 
clearances. As all these remnants are strongly influenced by grazing we do not know their 
natural state, but the presence of Abies spectabilis, Juniperus recurva, and Rhododendron spp. , 
partly covered with Usnea longissima, suggests that cloud forests may be the potential 
vegetation, as on the shady slopes. If these conclusions are admitted, it follows that the highest 
altitudinal records of trees show distinct differences on different slopes. The highest altitude at 
which I encountered tree-forming species was over 4400 m. The very highest record was a 
Juniperus recurva south of the Ngozumpa glacier at 4440 m. It was obviously a young specimen, 
60 cm tall and with a trunk diameter of 5 cm; the exposure was SSW. In the same valley and at the 
same exposure, I found several tree junipers between 4440 and 4420 m. The tallest were 3.5 m 
high with trunk diameters of 20 cm near the ground; all had been occasionally lopped. The 
highest grove of trees in the Khumbu Himal is situated around the houses of Pangpoche at 4240 
m, with a southerly exposure. The trees are 6 m high, the diameter at breast height is 25 cm at 
most, and the canopy is closed. It is obviously a holy grove, with prayer flags placed on trees 
whose branches have not been lopped. If these highest records indicate the minimum altitude of 
the upper limit of forests which could then be regarded as the climatic limit, it is obvious that the 
potential natural and climatic upper limit of forests in the valleys south of Mt Everest, especially 
in the Khumbu Himal, is at least 4400 m. This height for the potential upper limit of forest is 
clearly applicable only to the sunny slopes; on the shady slopes, it is at least 4200 m, which 
coincides with the highest records for individual trees. If the tree line is higher on the sunny than 
the shady slopes, it can be assumed that the more favourable temperature conditions of the 
sunny slopes determine the difference. Thus in these valleys at the northern borders of the 
tropics the upper Umit of forests is controlled by temperature. The rainfall is obviously not a 
limiting factor as an annual precipitation of as Uttle as 500 mm, concentrated in the growing 
period, is enough to make the sunny slopes the more favourable ones for forests. Thus the humid 
parts of the Himalayas and the European Alps have similar vegetational characteristics. 

This quasi theoretical upper forest Une is overlain by other factors in both mountain regions. 
In these south-eastern parts of the Himalayas, daily cloud formation in the late morning is 
characteristic, with the effect that the east-facing slopes get full sun during the first four or five 
hours of daylight , whereas the westerly slopes are under clouds in the afternoon , when otherwise 
the sun would warm this exposure. At first glance it seems quite obvious that this determines the 



232 G. MIEHE 

differences in the vegetation pattern of these valleys. However, there are only a few strictly east- 
and west-facing slopes and most are exposed to the south-east or north-east and, moreover, the 
south-eastern slopes are more easily accessible and more widely used for grazing and cultivation. 
Where strictly north-south running valleys occur, they are by contrast wooded on the eastern 
slopes and widely cleared on the west-facing slopes. 

Another factor overlying the influences of insolation is the valley wind which occur almost 
daily, starting up in the late morning. The exact effect of these winds has not yet been 
determined, but the difference between the vegetation on wind-exposed flanks compared with 
that found on lee slopes is striking. Slopes mainly covered by birch forests show firs on the 
exposed ridges, whereas fir-covered slopes have pines on the wind-blown parts. Near the upper 
limit of forest, where rhododendron woodlands become dominant, exposed slopes are devoid of 
trees, especially rhododendrons, but dwarf rhododendron cushions (R. setosum and R. antho- 
pogon) or dwarf prostrate junipers occur. It appears that on the wind-blown slopes, the 
vegetation is replaced by species that may be expected to be more drought resistant. As has been 
proved by experiments in the European Alps (Larcher, 1963), and with the knowledge that the 
growing season is undoubtedly humid while the winter is often dry and cold, one can assume that 
full solar radiation in winter, when the soil is frozen and water is not available, leads to 
transpiration rates that are significantly increased when the winds blow. This effect of seasonal 
drought due to freezing and enhanced by wind, probably leads to a tree-line which is naturally 
lower on the sunny and exposed slopes than on the shady lee slopes. As the leeward slope of a 
ravine in a north-south running valley is also the shady one, one can erroneously assume that the 
shady slope is more favourable for forests than the sunny slope. 

Remarks on the influence of grazing, burning, and wood-cutting 

In the area discussed there is great variation resulting from human influences. In the Barun 
Khola, the Iswa Khola, the Hunku Drangka, the Hinku Drangka, and the Lumding Khola, the 
cloud forests are in a condition which is fairly typical for the southern side of the main range. The 
upper cloud forest belt is cleared near the timberline for the seasonal settlements of summer 
grazing places. Burning is the common initial process in making clearings in order to improve the 
grazing conditions by the removal of the woody competitors of herbs. Usually the forests are 
penetrated by these clearings which are surrounded by areas of open woodland which have been 
gradually cleared by extensive uncontrolled grazing. 

In the alpine belt there are usually summer grazing places of transhumant sheep and goats. 
Here the herdsmen partly burn the moist alpine scrub consisting mainly of Rhododendron 
setosum. 

In the upper catchment of the Dudh Kosi, in the Khumbu Himal, the natural vegetation has 
been more extensively removed than is usual for this side of the Himalayas. Instead of seasonal 
settlements, the Sherpas have cleared the forests for the establishment of permanent settle- 
ments. Thus the influence of grazing is continuous throughout the year, in winter near the 
villages, in summer mainly in the lower alpine belt up to 5000 m. As the Khumbu Himal receives 
monsoon rains, there is relatively Uttle snow in winter (although heavy falls may occur). For this 
reason, only the calves are kept in the houses during the winter and most of the cattle graze 
around the villages. After snowfall, the cattle at first remain in the fields near the houses, but 
then concentrate on grazing the slopes where the snow melts first. The influence of grazing is 
strongest on the south-facing 30° slopes. Steepness of slope is not a major factor governing the 
amount of grazing and a 10° slope close to a village is usually less grazed than a 20° slope at some 
distance from a village. 

On these heavily grazed slopes, most of the palatable herbs other than grasses are displaced by 
inedible species like Iris spp. Euphorbia spp, species of Labiatae, Campanulaceae, Gentiana 
spp, orchids, and dense creeping Cotoneaster spp which stabilize the slopes. The duration of 
snow cover is a deciding factor in governing the type of vegetation. This explains partly why 
some forests occur on shady slopes close to the villages. Furthermore, these are to a certain 
extent protected by the indigenous rules of the Sherpas, who elect a forest warden whose duty is 



PLANTS SOUTH OF MT EVEREST 233 

to control wood-cutting, litter collecting, and even grazing in the forest (Furer-Haimendorf, 
1964). Thanks to these laws the Sherpas have as yet no shortage of wood for fuel. 

The fact that in the Khumbu Himal the natural vegetation is so largely replaced by the 
secondary associations is partly due to the relief. The south side of the main range is directly in 
contact with the Tibetan north slope and can be entered via the Nangpa-La, a pass which is only 
5716 m high and is passable even for cattle. The Sherpas, who originated on the eastern slope of 
the Tibetan Plateau, crossed the Nangpa-La in the first half of the sixteenth century (Oppitz, 
1968). It is quite probable that the original 20 to 50 settlers first practised shifting cultivation and 
cleared the sunny slopes by fire, while the cattle, preferring particular species such as Abies 
spectabilis, transformed the forests into open woodlands. Owing to the continually uncertain 
harvests the population remained small but, nevertheless, their influence on the vegetation was 
great because the burnt areas were invaded by Juniperus recurva. This species is very typical of 
the clearings (Stainton, 1972: 107) and, because it is not grazed by cattle, forms a comparatively 
stable secondary forest. Evidently there were larger forests of junipers in the Khumbu Himal in 
the nineteenth century. The potato was introduced into the Khumbu area about 1850 (Fiirer- 
Haimendorf, 1964: 8). A drastic population increase followed and the juniper forests have been 
increasingly used for fuel, so that these secondary forests have, in turn, become open 
woodlands. 

Alphabetical list of vascular plants, their life forms, associated formations, and 

records of collection 

Key to plant life form symbols 

The present information regarding plant life forms is based on the revised list of the Raunkiaer 
System of plant Hfe forms (EUenberg, 1956: 26) as completed by Ellenberg & Mueller-Dombois 
(1967). 

Phanerophytes (P) 
Mes P scap = large trees (5-50 m). 
Mi P scap = small trees (2-5 m). 
N P scap = dwarf trees (< 2 m). 
Mi P caesp = tall shrubs (2-5 m). 
N P caesp = normal-sized shrubs (< 2 m). 
N P gram = bamboos. 

Chamaephytes (Ch) 
Ch frut = woody dwarf-shrubs up to 50 cm, woodiness completed into branch-tips. 
Ch suff = semi- woody dwarf-shrubs. Woodiness restricted to the base of the shoot system. 
Ch herb = herbaceous chamaephytes: all non-woody perennial forbs, grasses, and ferns. Most of the 

non-rhizoid evergreen ferns of the Khumbu area should be included here, 
a = cold-deciduous (summergreen). 

e = evergreen, 

s = sclerophyllous. 

s/e, s/a = facultatively deciduous depending on altitude, shelter from frost, 
p = diphyWous {Ephedra). 

b = needle-leaved, 

rept = reptant. 
pulv = pulvinate. 

f pulv = flat cushions (Silene acaulis, Gentiana acaulis). 
g pulv = globose cushions {Androsace helvetica). 
1 succ = leaf succulents dying back to shoots, some are above the ground, mostly globose-cushions. 

Hemikryptophytes (H) 

Remnant shoots flat on the ground, often protected by dead shoot remains. Active shoots during growing 
season raised above the perennial ground-shoots. Typically herbaceous, the maturing stem may be 
lignificated. 
H caesp = branched or circular shoot system. 



234 G. MIEHE 

H rept = creeping and/or matting. 

Hscap = scapose without rosette. 

Hros = rosette. 

H sem = rosette, but with leaves on the stalk. 

H scap = without rosette, mainly tall forbs. 

Geophytes (G) 

Survival organs protected in the soil. 

G rad = root-budding geophytes. 

G bulb = bulbous geophytes. 

G rhiz = rhizome geophytes. 
With some species information concerning the supraterrestrial growth habit is given (caesp, scap, sem, 
rept . . .). 

Therophytes (T) \^ 

Live less than a year. Growth habit see hemikryptophytes. 

Lianas (L) 

PL = phanerophytic and chamaephytic lianas including all climbing plants that do not die back 

periodically. There are woody (frut), semi-woody (suff), and herbaceous (herb) life forms. 
rPL = root climbers. 
stPL = winding climbers. 
elPL = tendril climbers. 
dPL = spread climbers. 

HL = hemikryptophytic climbers, dying periodically to a remnant shoot system near the ground. 
GL = geophytic lianas, dying periodically to subterranean storage organs. 
TL = therophytic lianas. 

Epiphytes (E) 
fac E = some phanerophytic, mainly chamaephytic woody or semi-woody facultative 

epiphytes, creeping on moss-covered trunks or boulders in the cloud forest belt, 
obi Ch E succ = obligative epiphytes with strongly modified root system and succulent leaves. 
GE bulb = bulbophytic, in moss of tree trunks. 

Further morphological characteristics 

rept/sem : facultatively straggling, creeping. 

G rad/H sem : with storage root, sometimes shoot under the surface, sometimes near the surface. 

Par : parasitic. 

hyd : water plant. 

fac hyd : facultatively in water (flushes, wet rocks). 

In the free solifluction belt changing life forms are characteristic: without the daily moving solifluction scree 
most of the high altitude phanerogams would be of the hemikryptophytic or flat cushion habit. So, in flat 
sites, where the scree movement is little, the genetically fixed life form is typical, whereas on solifluction 
slopes the shoots are deformed by adapting to this movement and the plants become rhizoid. 

* Drawing or colour photograph in Polumin & Stainton (1984). 

-I- Not included in Hara, Stearn & Williams (1978), Hara & WiUiams (1979), and Hara, Chater & 
Williams (1982). 

Key to plant formation symbols 

Cloud forest belt 

I cloud forest (medium belt) between 2400 m/8000 ft and 3200 m/10500 ft which is altitudinally 
characterized by the huge Tsuga dumosa trees; tree-trunks and branches are moss-covered. 

II open, mostly south-facing, wind-exposed grazed pine forests {Pinus wallichiana) up to 3500 m/11 500 
ft. 

III cloud forest (upper belt) between 3200 m/10500 ft and the upper tree line (4200 m/13800 ft) with 
Abies spectabilis, Betula utilis, and Rhododendron thickets. Beard lichens {Usnea longissima) are 
characteristic. 



PLANTS SOUTH OF MT EVEREST 235 

IV wet, moss-covered rocks in the forest belt with species of mainly subtropical distribution (Corallodis- 
cus, Spathoglottis) and alpine cushions (Rhodiola, Saxifraga). 

V grazing grounds around the villages, with the plants originating from forest clearings and forest 
undergrowth, and alpine plants spreading down to the potential forest belt after deforestation. 

Alpine belt 

VI mostly south-facing, wind-blown dwarf juniper scrublands between 4000 m/13200 ft and 5000 
m/16500ft. 

VII mostly north-facing, wind-sheltered moist alpine scrub {Rhododendron anthopogon, R. setosum, 
R. nivale) between 3800 m/12500 ft and 5200 m/17000 ft. 

VIII flushes. 

IX Kobresia pygmaea - mats between 5000 m/16 500 ft and 5500 m/18 000 ft. 
Free solifluction belt 

X phanerogams adapted to the soHfluction moved scree up to the highest open slopes at about 6000 
m/19700ft. 

Abbreviation list of collectors' names 

The cited specimens are mainly those of the BM. The plants collected by A. Byers, J. F. 
Dobremez, J. Poelt, G. F. Smith, L. W. Swan, T. Wraber, and A. Zimmermann are partly cited 
according to duplicates in BM or determination lists which I was able to consult by the kind 
support of Susan Sutton (BM), Annelies Schreiber (M), Alton Byers (COLO), and George F. 
Smith (Stockport). Collections made after 1979 are mentioned by the date and partly by the 
number of collected plants and the herbarium. For further information and the collections 
before 1979 see Sutton (1978) and Hara, Chater & Williams (1982: 8). 

B = M. L. Banerjee. Cited specimens in BM. 

Beer = L. Beer. Cited specimens in BM. 

BL = S. Bowes Lyon. Cited specimens in BM. 

BLM = L. Beer, C. R. Lancaster, D. Morris. 

By = A. Byers. Between March and November 1984, c. 2500 specimens. Cited according to the 

collector's list. The collection will be given to COLO. 
D = J. F. Dobremez. Cited specimens in BM and according to the collector's list (BM). 

Einarss = S. Einarssen. Cited specimen in BM. 

dH. = J. de Haas. Between August and September 1974. Cited specimens in BM. 
H = A. Horsfall. Cited specimens in BM. 

Heub. = H. Heuberger, pers. comm. Record oiArenaria bryophylla between 6000 and 6100 m, Cho 

Oyu NE ridge, 1954. 
L = Lall Dhwoj . Cited specimens in BM. 

Mc = D. J. McCosh. Cited specimens in BM. 

M = G. Miehe. Between August and November 1982, 318 specimens, in the BM. 

N = D. H. Nicholson. Cited specimens in BM. 

P = J. Poelt. Cited specimens in BM and according to the collection list compiled by A. Schreiber 

(M). 
S = J. D. A. Stainton. Cited specimens in BM. 

Sch = A. T. SchiUing. Cited specimens in BM and according to collector's list (BM). 
Sh = Puspa Ratna Shakya. Cited specimens in BM. 

Sharma = Khadanand Sharma. Cited specimen in BM. 
Sm = G. F. Smith. Between July and August 1984, above 4500 m. Cited specimens according to 

'provisional list of plants . . .', typescript, Stockport, Gottingen. 
Sw = L. W. Swan. Cited specimens in BM and according to collector's list (BM). 
Tilman = H. W. Tilman. Cited specimen in BM. 

W = T. Wraber. Cited specimens in BM and according to determination Ust (BM). 

Z = A. Zimmermann. Cited specimens according to duplicates in BM and according to CandoUea 

15-17. 

Alphabetical list of flowering plants 

The plant names and their authors follow y4rt enumeration of the flowering plants of Nepal (Hara, 
Stearn & Williams, 1978; Hara & Williams, 1979; Hara, Chater & Williams, 1982). 



236 

Abies spectabilis (D. Don) Mirb. ' 
Acanthocalyx nepalensis (D. 
Don) M.J. Cannon * 



Acanthopanax cissifolius (Griff. 

ex Seem.) Harms * 
Acer caudatumV^dW. * 



A. pectinatum Wall, ex Pax * 
A. stachyophyllum Hiern 
Aconitum balfourii Stapf 
A. ferox Wall, ex Seringe * 

A. heterophyllum Wall, ex Royle 
A. hooker i Stapf * 



A. laciniatum (Briihl) Stapf var. 

laciniatum 
A. rotundifolium Kar. & Kir. 
A. spicatum (Bruhl) Stapf * 
Aconogonum campanulatum 

(Hook.f.) Hara var. 

campanulatum * 
A. campanulatum var. oblongum 

(Meisn.) Hara * 
A. molle (D. Don) Hara 
A. rumicifolium (Royle ex Bab.) 

Hara* 
A. tortuosum (D. Don) Hara * 

Acronema cf. hookeri (C. B. 

Clarke) H. Wolff 
A. johrianum Babu + 
A. tenerum (DC.) Edgew. 
Aechmanthera gossypina (Wall.) 

Nees * 
Aeschynanthus hookeri C. B. 

Clarke 
Agrostis munroana Aitch. & 

Hemsley 
A. nervosa Nees ex Trin. 
A. pilosula var. wallichiana 

(Hook.f.) Bor 
A. triaristata (Hook.f.) Bor. 
Ainsliaea aptera DC. * 
Ajuga bracteosa Wall, ex Benth. 
A. lobata D. Don * 
Aletris pauciflora (Klotzsch) 

Hand.-Mazz. * 
Allium wallichii Kunth * 
Alnus nepalensis D. Don * 
Anaphalis busua (Buch.-Ham. ex 

D.Don) DC. 
A. cavei Chatterjee 
A. contorta (D. Don) Hook.f. 



G. MIEHE 






b Mes P scap 


III 


Sch.2342, W.436 ■> -^^ 


aHsem 


V,VI 


By.l2,D.337,380, 
BL.2058,2135,M.903, 
Mc.322, Z.694, 1474, 
Sm.s.n. 


a NP caesp 


I 


BL.2090, Z.446, 760 


a Mi P scap 


III 


BL.2172,By.47,326,640, 
dH.2967, Sch.2348, 
2349,2261,2262 


a Mi P scap 


I 


S.2246, 2247, 5946 


a Mi P scap 


I 


Sch.2361 


a G bulb 


V 


Z.1796 


a G bulb 


i,v 


dH.2900,S.6577,P.s.n.,P 




V 


B.78, Z.1309, 1661 


a G bulb 


Z.1796 A 


a G bulb 


V-VII 


By.ll32,H.50,M.1009, 
P.s.n.,S.6595,W.196, ' 
Sm.s.n. 


a G bulb 


V,VI 


S.6577 


a G bulb 


I 


P.s.n. 


a G bulb 


V 


M.875,P.s.n.,W.289 


aGrad 


I 


Z.1889 



aGrad 



Z.787 



aGrad 
a G rhiz 


II, III 
VII 


M.845,Z.1691A 
D.404, P.s.n. 


aGrad 
aGrad 


VI 
III,V 


D.340,dH.1970,L.0656, 

Sm.s.n., Sch.2040,Z.66'/ 
P.R.162 


aGrad 
a Ch suff 


? 
VII 

I 


By.993 

W.282 
P.69 


sobl.dEPL 


I 


S.4488 


a H caesp 


VI 


P.s.n. 


a H caesp 
a H caesp 


VI 

V 


M.892b, W.366a 
W.429 


a H caesp 
aHros 
a H caesp 
a H rept 
aGrad 

aGrad 

a/e Mes P scap 

a H scap/Ch suff 


V 

VI 
VI 

I 

VI 

VII, VIII 

I 

I 


W.350 

Z.413 

By. 893 

Z.774 

BL.2043, By.73, D.334, 

397, Z.621, 669, Sm.s.n. 
By.625,634,680,W.409 
Z.1868 
f.M. 


a f Ch herb pulv 
aHsem 


VI, IX 
V 


M.1037, Sm.s.n. 
By.169,314,385,418,742, 
H.3, P.G.298 





PLANTS SOUTH OF MT EVEREST 


237 


A. margaritacea (L.) Benth. * 


aHsem 


n 


By.370, dH.2879 


A. royleana DC. 


a H rept/sem 


n 


W.303 


A. tenella DC. 


a H rept/sem 


V 


D.384,dH.2917,Z.1755 


A. tripUnervis (Sims) C. B. 


aHsem 


V 


By.411,1130,D.332, 


Clarke * 






Z.1089 


A.triplinervis vzx . intermedia 


aHsem 


V 


H.67, ?Z.s.n. 


(DC.) Airy Shaw 








A. tripUnervis var. monocephala 


a f Ch herb pulv/a 


VI 


D.343,M.1015,Sm.s.n. 


(DC.) Airy Shaw* 


Hsem 






A. xylorhiza Sch.Bip. ex Hook.f. 


a f Ch herb pulv 


VI, VI, IX 


H.47,BL.2110,D.414, 
W.210,Sm.s.n. 


Andropogon munroi C. B. Clarke 


a H caesp 


IV 


M.817b, Z.1733 


Androsace delavayi Franch. * 


e f Ch herb pulv 


IX, X 


BL.2123,Sm.s.n. 


A. geraniifolia Watt 


aHros 


I, IV 


S.4592 


A. globifera Duby * 


e f Ch herb pulv 


IX 


D.416,Mc.323,M.1045, 
S.637 


A. hookeriana Klatt 


aHros 


VIII 


Z.823 


A. lehmannii Wall, ex Duby * 


e g Ch herb pulv 


IX 


BL.2008, D.390, Mc.284, 
Z.816 


A. nortonii Ludlow 


aHros 


IX 


BL.2046,By.445,D.412, 
Mc.318,Sm.s.n. 


A. sarmentosa Wall. * 


aHros 


II, V 


By.46, 227, 1092, dH.2923, 
Z.466, 686 


Anemone demissa Hook.f. & 


aHsem 


V,VII 


BL.2109,D.423,Z.601, 


Thorns. * 






693, Sm.s.n. 


A. obtusiloba D. Don* 


aHsem 


III 


P.R.306 


A. rivularis Buch.-Ham. ex DC. * 


aHsem 


i,v 


By.393, M.902, P.s.n. 


A. rupestris Wall ex Hook.f. & 


a G bulb 


I, III 


BL.2087 


Thorns. 








A. rupicola Cambess. * 


a G rhiz 


I, III, VI, VII, 
VIII 


S.669 


Arabidopsis himalaica (Edgew.) 


Tsem 


III, V, IX 


By.l21,W.297 


O. E. Schulz 








A. lasiocarpa (Hook.f. & 


T caesp 


i,v 


B.&Sh. 5656 


Thorns.) O. E. Schulz 








Arctium lappa L. 


a H caesp 


V 


f.M. 


Arcyosperma primulifolium 


aHros 


VII 


Mc.52 


(Thorns.) O. E. Schulz 








Arenaria bryophylla Fernald * 


a g Ch herb pulv 


VI,IX,X 


H.33,M.1076, P.s.n., f. 
Heub. 


A. ciliolata Edgew. 


a H rept/ros 


VIII, X 


H.41,M.1043, Sm.s.n. 


A. debilis Hook.f. ex Edgew. & 


a H caesp 


V 


By. 637 


Hook.f. 








A. densissima Wall, ex Edgew. & 


a g Ch herb pulv 


IX 


By.797, D.410, Mc.2038, 


Hook.f. * 






S.7115 


A.edgeworthiana Majumdar 


a g Ch herb pulv 


VI, IX 


Mc.377, W.227, Sm.s.n. 


A. festucoides Benth. 


a g Ch herb pulv 


IX, X 


Sw.s.n. 


A. glanduligera Edgew. ex 


a f Ch herb pulv 


VIII, X 


By.385,H.28,M.955, 


Edgew. & Hook.f. * 






Mc.377, P.s.n., W.178, 
231, Sm.s.n. 


A. glanduligera var. cernua N. F. 


a f Ch herb pulv 


VIII, X 


M.1074 


Wilson 








A. globiflora Edgew. & Hook.f. 


a g Ch herb pulv 


VI 


BL.2073, Mc.309, S.7095 


A. melandryoides Edgew. ex 


a G rad/rept 


IX 


W.189, Sm.s.n. 


Edgew. & Hook.f. 








A. polytrichoides Edgew. ex 


a g Ch herb pulv 


IX 


M.989, S.7157 


Edgew. & Hook.f. 








A. pulvinata Edgew. ex Edgew. 


a f Ch herb pulv 


IX 


By.773, Sw.s.n. 



& Hook.f. 



238 



G. MIEHE 



Arisaema jacquemontii Blume * 
A. speciosum var. mirabile 

(Schott) Hara 
A. vexillatum Hara & Ohashi 
Artemisia biennis Willd. 
A. capillaris Thunb. 
A. dubia Wall, ex Besser * 
A. indica Willd. 
A. striata Edgew. 

A. wallichiana Besser * 
Aruncus dioicus (Walter) Fernald 
Arundinaria mating Gamble 
Aster albescens (DC.) 

Hand.-Mazz. * 
A. barbellatus Grierson 
A. diplostephioides (DC.) C. B. 

Clarke * 

A. flaccidus Bunge * 

A. himalaicus C. B. Clarke * 

A. sikkimensis Hook. 

A. stracheyi Hook.f. 

A. trinervius Roxb. ex D. Don * 
Astilbe rivularis Buch.-Ham. ex 

D. Don * 
Astragalus chlorostachys Lindl. 
A. concretus Benth. 
A. donianus DC. 

A. floridus Benth. * 
A. frigidus (L.) A. Gray 

A. strictus Grab, ex Benth. 

Begonia picta Smith * 
Berberis angulosa Wall, ex 
Hook.f. (feThoms.* 

B. ceratophylla G. Don 

B. concinna Hook.f. 

B. cf. erythroclada Ahrendt 

B. jaeschkeana C. K. Schneid. 

B. mucrifolia Ahrendt 

B. tsarica Ahrendt 

B. virescens Hook.f. 

Bergenia purpurascens (Hook.f. 

«feThoms.)Engl. 
Betula alnoides Buch.-Ham. ex. 

D. Don * 
5. uri/wD.Don* 

Bistorta affinis (D. Don) Greene 

B. amplexicaulis (D. Don) 
Greene * 



a G bulb 


I 


BL.2064 ^ 


a G bulb 


I,III,V 


BL.2029 


a G bulb 


I 


BL.2055 


a H scap 


II, V 


L.0181,P.G.268 


a H scap 


V 


Z.1792 


a H scap 


V 


Z.1655 


a H scap 


V 


P.s.n. 


a H scap 


V 


M.836,P.s.n.,W.200, 
Z.1792 


a H scap 


VIII 


B.5707, dH.2997, S.6589 


a Ch suff 


III 


By.297, 464, Beer 25447 


a/e NP gram 


I, III 


W.347 


a H sem/Ch suff 


II 


M.828,Mc.392,W.361, 
Z. 1635, 1653 


a G rhiz/sem 


V, VII 


D.386,415 


aHsem 


V,VI 


By.696, dH.2953a, M.899, 
P.R.298,Z.1717, 1822, 
Sm.s.n. 


a G rhiz/sem 


VIII 


BL.2128,D.415,Sm.s.n. 


a G rhiz 


VI, IX 


By.ll69,M.869, 1042, 
Sch.2173 


a H sem/Ch suff 


I 


Sch.2210, 2250, Z. 1880, 
1885 


a G rhiz/rept 


VIII 


Beer 25388, By.564, 
Mc.313 


a H sem/Ch suff 


I 


Z. 1880a 


aGrad 


I 


P.44 


a H scap 


II, III 


By.321,Z.1768A 


a H rept/caesp 


V 


BL.2155, S.7100 


a H rept 


V,VI 


BL.2132,By.210,311,312, 
339 


a H scap 


V 


S.7152 


a H caesp/rept 


V 


P.s.n. 


s Ch frut rept 


V,VI 


M.1013,Mc.314 


a/e G bulb fac E 


I, IV 


P.77 


a NP caesp 


V 


Sch.2270, Z.494, 1619 \ 


a/e NP caesp 


II, V 


Z.1835 


a/s NP caesp 


V 


By.l012 


a/e NP caesp 


V 


M.992 


a/e NP caesp 


V 


By.717, 720, 967, 1112, 
1154,Z.1758 


a Ch frut 


V,VI 


BL.2103 


a NP caesp 


V 


Z.593 '" 


a/e NP caesp 


II, V 


Z.480 


a G rhiz/ros 


IV 


By.l75,Sch.2323,Z.653 


a Mes P scap 


I 


f.M. 


a Mes P scap 


III 


By.ll57,W.434,Z.406, 
492 


a G rhiz 


VI, VII 


By.478,D.365,Z.1481, 
Z.1573, Sm.s.n. 


aGrad 


V,VII 


By.l53,211,M.849,913, 



P.G. 348, Sch.2344 



B. macrophylla D. Don) Sojlk 



PLANTS SOUTH OF MT EVEREST 
a G rad/sem VI, VII, IX 



239 



BL.2198, M.948, Mc.330, 
P.s.n.,S.4521,W.201, 
279, Z.737, 1478, 
Sm.s.n. 



B. milletii Lev. 


aGrad 


V 


W.s.n.,Z.1478 


B. perpusilla (Hook.f.) Greene 


a G rad/rhiz 


VI, IX 


Beer 9552 


B. vaccinifolia (Wall, ex Meisn.) 


a Ch frut rept 


III 


By.75,82, 137, 174,348, 


Greene * 






502,M.861,Sch.2346, 
2175, 2181, Z.609, 1709 


B. vivipara (L.) S. F. Gray 


a G rad/sem 


VI, VII, IX 


D.366, 391, Z.640, Sm.s.n 


Biswarea tonglensis (C. B. 


stTL 


I 


S.7190, Z.785 


Clarke) Cogn. 








Boschniaka himalaica Hook. & 


a G bulb 


V 


Mc.306 


Thorns, ex Hook.f. * 








Brachyactis anomala (DC.) 


aHsem 


V 


M.911 


Kitam. 








Bray a oxycarpa Hook.f. & 


a G rad/H caesp/f 


X 


Z.564, 597 


Thorns. * 


Ch herb pulv 






Buddleja crispa Benth. * 


a NP caesp 


I 


Z.262 


Bulbophyllum retusiusculum 


obi e E Ch succ 


I 


L.0433 


Reichenb.f. 








Bupleurum candollii Wall, ex 


aGrad 


I 


D.322, P.72 


DC* 








Cacalia pentaloba Hand.-Mazz. 


a H scap 


III 


B.5651,M.978 


Calamagrostis emodensis Griseb. 


a H caesp 


? 


W.394 


C epigejos (L.) Roth 


a H caesp 


III,V 


Z.1771 


C. pseudophragmites (Hall.f.) 


a H caesp 


VIII 


Sm.s.n. 


Koel. ex. Baumg. 








Calanthe alpina Hook.f. ex Lindl. 


aGrad 


I 


Mc.265 


Callitriche palustris L. 


a hyd H caesp 


IV 


M.1114,P.s.n. 


Campanula aristata Wall. 


aHsem 


V-VII 


By.427, D.462, Mc.361 


C. modesta Hook.f. & Thorns. * 


a H scap 


V 


By.549,581,L.2138, 
Sm.s.n. 


C. nafcaoi Kitam. 


a G rhiz 


VI, VII 


S.1666,W.295 


C. pallida Wall. 


a H scap 


V 


By.307,315,Z.1654 


Capsella bursa-pastoris (L.) 


Tsem 


V 


By.214, P.78, Z.418, 1764 


Medikus 








Cardamine inayatii O. E. Schulz 

C. loxostemonoides O. E. Schulz 

* 


T caesp 


I, II 


BL.2016 


a H/T scap 


V 


BL.2111,Mc.360 


C. scutata suhsp.flexuosa 


Tscap 


I, II 


Z.788 


(Withering) Hara 








C. trifoliolata Hook.f. & Thorns. 


Tscap 


1,11 


Mc.272 


Cardiocrinum giganteum (Wall.) 


a G bulb 


I 


P.s.n.,Sch.2248,Z.1898 


Makino * 








Carex atrata subsp. pullata 


a G rhiz 


VIII 


BL.2166,D.446 


(Boott) Kiikenthal 








C. atrofusca Schkuhr 


a G rhiz 


VIII 


D.376,M.1094,P.G.288, 
R.331 


C. inanis Kunth 


a G rhiz/caesp 


IV, VI, VIII 


BL.2160 


C. infuscata Nees 


a G rhiz 


VI, VII 


M.958, 1014 


C. laeta Boott 


a H caesp 


VI, VIII 


P.R.338, Z.696 


C. lehmannii Drejer 


a G rhiz 


VIII 


P.R.121,R.325 


C munda Boott 


a G rhiz/caesp 


VIII 


W.310 


C. nakaoana T. Koyama 


a H caesp 


VIII 


dH.2994,M.1029,P. 
G.219, G.292, G.297, 
S.666,W.169 



240 


G. MIEHE 






Carpesium nepalense Less. 


a H scap 


IE 


W.397 


Cassiope fastigiata (Wall.) D. 


s Ch frut 


vn 


By.469, 687, D.379, 


Don* 






W.248, Z.612, 634, 702, 
1559, 1756, Sm.s.n. 


Castanopsis tribuloides (Sm.) A. 

DC* 
Celastrus stylosus Wall. 


s Mes P scap 


I 


f.M. 


a d LP frut 


I 


S.4484 


Cerastium fontanum Bautng. 


a H rept 


? 


By.498 


Chamaesium novemjugum (C B. 


aGrad 


VI, IX 


M.936 


Clarke) C Norman 








Chenopodium botrys L. 


Tscap 


V 


By.724, P.s.n. 


C. graveolens Willd. -f 




? 


By. 1124 


Chesneya nubigena (D. Don) AH 


a f Ch frut pulv 


IX 


BL.2112,By.41,M.1035, 


* 






Mc.329, W.249, 
Sm.s.n. 


Chionocharis hookeri (C B. 


e g Ch herb pulv 


X 


Sw.224, 225 


Clarke) J. M. Johnston * 








Chrysosplenium alternifolium 


H caesp/scap 


IV 


Z.805 


var. sibiricum Seringe ex DC. 








C. carnosum Hook.f. & Thorns. * 


H caesp 


III 


By.38,S.7117 


C. cf. griffithii Hook.f. & Thorns. 


a G rhiz/scap 


III 


M.983 


C. lanuginosum Hook.f. & 


a G rhiz/sem 


I, III 


Z.458 


Thorns. 








C. nudicaule var. intermedium 


G rhiz/scap 


VIII 


BL.2035, Mc.53 


Hara 








Cicerbita cyanea (D. Don) 


a G rad/H scap 


II 


dH.2950a, P.s.n., Z. 1791 


Beauv. 








C. macrantha (C B. Clarke) 


a Grad/H scap 


V 


P.s.n. 


Beauv. * 






(■ 


C. macrorhiza (Royle) Beauv. * 


aGrad 


II, V 


By.586, 712, P.s.n. 


C. macrorhiza var. saxatilis 


a G rad/a H scap 


IV, V 


dH.2941,M.128 


(Edgew.) Beauv. 








Circaea alpina subsp. micrantha 


T rept/scap 


? 


W.317 


(Skvortsov) Bufford + 








C. alpina subsp. imaicola 


T rept/scap 


II 


P.7 


(Aschers. & Magnus) Kitam. 








C. repens Wall, ex Aschers. & 


T rept/scap 


I, IV 


S.5750 


Magnus 








Cirsium falconeri (Hook.f.) 


a G rad/sem 


V 


W.377,Z.1670 


Petrak * 








C. verutum (D. Don) Spreng. * 


a G rad/scap 


V 


By.601 


Clematis montana Buch.-Ham. ex 


a el PL frut 


ni 


By. 654, dH.2975, 


DC* 






Sch.2186, Z.502 


Clinopodium umbrosum (M. 


a H caesp 


in 


P.s.n., W.307,Z. 1670 


Bieb.) C.Koch 








Clintonia udensis var. alpina 


a G rhiz 


m 


By.26, Z.641 


(Kunth ex Baker) Hara * 








Coccinea grandis (L.) Voigt 


TL 


I 


P.s.n. 


Codonopsis ovata Benth. * 


a H scap 


V 


By.362 


C. thalictrifolia Wall. 


a G rad/rept 


V-VII 


By.342, 430, 453, M. 1058, 
Sm.s.n. 


Coleus barbatus (Andrews) 


a H scap/Ch suff 


II 


Z. 1833, 1865 


Benth. * 








Colutea multiflora Shap. * 


a NP caesp 


i,n 


dH.2951, Sch.2165 


Commelina paludosa Blume * 


a H scap 


n 


By.465 


Corallodiscus lanuginosus (Wall. 


eHros 


IV 


D.321,Mc.394,M.811, 


ex DC.) Burtt * 






Z.422, 1225 


Cortia depressa (D. Don) C 


aGrad 


VI, IX 


BL.2179,D.368, P.s.n., 


Norman * 






G.259, Sm.s.n. 



Cortiella hookeri (C. B. Clarke) 

C. Norman * 
Corydalis cashmeriana Royle * 
C. casimiriana Prain 
C. caveiT). G. Long 
C. chaerophylla DC. 
C. chasmophila Ludlow 
C. flabellata Edgew. 
C. flaccida Hook, f .& Thorns. 
C. govaniana Wall. * 
C. hookeri Prain 
C. /Mncea Wall. * 
C. leptocarpa Hook.f. & Thorns. 
C. longipes DC. 

C. meifolia Wall. var. meifolia * 

C. polygalina Hook.f. & Thorns. 
C. stracheyi Duthie ex Prain + 
Corylus feroxV^^W. * 
Cotoneaster acuminatus Lindl. 

C. adpressus Bois 

C. /ngidM.s Wall, ex Lindl. * 

C. integrif alius (Roxb.) Klotz 

C. meuselii Klotz 

C. microphyllus Wall, ex Lindl. * 

C. nitidus Jacques 

C. sanguineus Yii 

Cremanthodium decaisnei C. B. 

Clarke * 
C. e//w« (Hook.f.) Kitam. * 

C. nepalense Kitam. * 

C. oblongatum C. B. Clarke * 

C. reniforme (DC.) Benth. * 

C. rerM.sww (Wall, ex Hook.f.) R. 

Good* 
Crepis tibetica Babcock 
Cryptothladia polyphylla (Wall. 

exDC.)M. J. Cannon* 
Cuscuta reflexa Roxb. 
Cyananthus hookeri C. B. Clarke 

C. incanus Hook.f. & Thoms. * 



C. inflatus Hook.f. & Thoms. 

C. lobatus Wall, ex Benth. * 
C. microphyllus Edgew. * 
C. pedunculatus C. B. Clarke 
C. spathulifolius Nannfeldt 
Cyanotic vaga (Lour.) J. A. & J. 

H. Schultes * 
Cymbidium hookerianum 

Reichenb. f. * 



PLANTS SOUTH OF MT EVEREST 


aGrad 


IX, X 


a G bulb 


V,VII 




V 




V 




I 


a H caesp 


vn 




? 




Ill 


a H caesp 


III,V 




VII 


a G bulb/scap 


III,V 




III 


a G rad/caesp 


III 


a H caesp 


VII 


a G bulb 


vn 


a H caesp 


VI 


a Mes P scap 


I 


a N P caesp 


II, V 


a Ch frut rept 


II, V 


a Mi P caesp 


II 


s Ch frut rept 


II, V 


a N P caesp 


II 


s Ch frut rept 


V 


s/a N P caesp 


II, V 


a N P caesp 


II 


aHros 


VIII, X 


aHros 


VIII, X 


aHros 


VII 


aHros 


VII, X 


aHsem 


vn 


aHsem 


vn 


a H ros/sem 


? 


aGrad 


V,VII 


ELTPar 


II 


a H rept 


V,VI 


a G rad/H rept 


V,VI 


a H rept/scap 


V 


a G rad/H rept 


V 


a G rad/H rept 


V 


a H rept 


V,VI 


a G rad/rept 


VI, VII 


aH 


n 


fac. e Ch E succ 


I 



241 

M.1001,W.164,233, 

Sm.s.n. 
D.431,Mc.283,P.s.n. 
H.lO(f.Liden) 
H.10(f. Liden) 
Mc.266 

BL.2163, S.628 
By. 193 

Mc.295, S.8360 
M.879, Z.809, Sm.s.n. 
S.7153 

BL.2164,Mc.341,S.4538 
S.4536 
By.525, 1167, dH.2935, 

M.850, W.386 
M. 1083, Beer 9555, 

Sm.s.n. 
Mc.327 
D.357 
S.4606 
dH.2930, M.854, Mc.273, 

S.4614, Sch.2030 
Sch.2031,2263 
Sch.2442A 
S.7125 
Sch.2355 

By.l089,P.s.n.,Sch.2032 
M.932 

Mc.304, BL.2168 
BL.2121,Mc.371, Sm.s.n. 

M.1108,W.165,Z.1456B, 

Sm.s.n. 
By.592,H.46, Sm.s.n. 
By.545,M.901,966, 

Z.1471 
By.506, 639, M.895, 

Z.1451 
Z.1451 

W.299 
Mc.381 

P.s.n. 
H.17,53,M.831,915, 

W.206 
By.76, 299, 328, 678, 1082, 

dH.2943, W.238, 

Z.1648, 1714, Sm.s.n. 
By.492,H.9,M.914,P.85, 

88 
P.89 

H.5, 55, P.94 

By.587, 795, M.873, P.s.n. 
Mc.308 
By.466 

S.4476 



242 

Cynanchum auriculatum Wight 
Cynoglossum glochidiatum Wall, 
ex Benth. * 

C. zeylanicum (Vahl) Thunb. ex 
Lehm. 

Cypripedium himalaicum Rolfe 
apud Hemsl. * 

Dactylorhiza hatagirea (D. Don) 

So6* 
Danthonia cachemyriana Jaub. & 

Spach 

D. cumminsii Hook.f. 
Daphne bholua Buch.-Ham. ex 

D.Don 
D. retusa Hemsl. * 
Datura stramonium h. * 
Delphinium caeruleum Jacquem. 

exCambess. 
D. drepanocentrum (Briihl) Munz 

D. glaciate Hook.f. & Thorns. 

D. cf. nepalense Kitam. & 

Tamura 
D. vwco5um Hook.f. &Thoms. * 
Desideria nepalensis Hara 
Desmodium elegans DC. * 
D. heterophyllum (Willd.) DC. 
Deutzia bhutanensis Zaikonn. * 
D. compacta Craib * 
D. staminea R. Br. ex Wall. * 
Deyeuxia pulchella (Griseb.) 

Hook.f. 
Dicentra scandens (D. Don) 

Walp. 
Dichrocephala benthamii C. B. 

Clarke 
D. integrifolia (L.f.) O. Kuntze 
Didymocarpus aromaticus Wall. 

ex D. Don 
D. oblongus Wall, ex D. Don 
D. primulifolius , D. Don * 
Diphylax urceolata (C. B. Clarke) 

Hook.f. 
Diplarche multiflora Hook.f. & 

Thoms. 
Dipsacus inermis var. mitis (D. 

Don) Y. Nasir 
Disporum cantoniense (Lour.) 

Merr. * 
D. cantoniense var. parviflorum 

(Wall.) Hara 
Dodecadenia grandiflora Nees * 
Draba altaica (C. A. Meyer) 

Bunge * 
D. amoena O. E. Schulz * 
D. elata Hook.f. & Thoms. 



G. MIEHE 






a d PL frut 
a H scap 


I 
II, V 


Mc.395 
B.5659,Mc.275,P.193 


a H scap/caesp 


II, V 


By.397, 872 


a G rhiz 


VII 


BL.2199,By.l20,D.395, 
Sm.s.n. 


aGrad 


V 


By. 142 


a H caesp 


V 


M.892C, P.s.n. 


a H caesp 
s NP caesp 


I 
I 


M.877, 851b, W.349 
S.4480 


s NP caesp 
a H scap 
aHsem 


I 

V 
II, V 


W.328 

f.M. 

P.s.n. 


aHsem 
aHsem 
aHsem 


V 
X 

VII 


By.423,537,621,dH.2969, 

L.0191,M.882 
M.1060, P.s.n., W.172, 

219, Sm.s.n. 
M.1033 


aHsem 

a NP caesp 
a NP caesp 
a NP caesp 
a NP caesp 
a NP caesp 
a G rhiz 

adHL 


VII 
X 

n 
n 
n 
n 
n 

III, V-VIII 

n 


Beer 25387, W.372 

Sw.71-72 

By.364 

P.s.n. 

Mc.38, S.4621 

BL.2157, By.499, S.7123 

Sch.2087, Z.472, 1860, s.n. 

M.817, 817a, 1021b, 1028, 

W.247 
P.s.n., Z. 1883 


aHsem 


I, II 


Z.749B 


aHsem 

a G bulb/scap 


n 

IV 


Z.749 
Z.746 


a G bulb/scap 
a G bulb/scap 
aGrad 


IV 
IV 
IV 


S.7163 
S.7164 
L.0529 


s Ch frut/rept 


III 


Beer 25361, Sw.441 


a H sem/scap 


1,111 


Z.1671 ' 


a G rad/scap 


II 


Z.765 


aGrad 


II 


Sch.2366A 


s Mes P scap 

a H ros/a f Ch herb 

pulv 
aHsem 
aHsem 


I 

X 

V 

VII, VIII 


S.6559 
Mc.325,W.167 

BL.2178, D.424. S.7144 
BL.2114, D.440, Mc.287, 
Z.521,581 



PLANTS SOUTH OF MT EVEREST 



243 



D. ellipsoidea Hook.f. & Thorns. 
D. gracillima Hook.f. & Thorns. 
D. lasiophylla Royle 

D. lasiophylla var. leiocarpa 

(Pamp.) O. E. Schulz 
D. cf. oariocarpa O. E. Schulz 

D. oreades Schrenk * 

Drosera peltata Smith 
Drymaria cor data (L.) Willd. ex 

Roem. & Schuh. 
Dubyaea hispida DC. * 



Elaeagnus infundibularis 

Momiyama 
Elsholtzia concinna Vautier 
E. densa Benth. 
E. eriostachya (Benth.) Benth. 

E. eriostachya var. pusilla 

(Benth.) Hook.f. 
E. fruticosa (D. Don) Rehder ' 



E. strobilifera (Benth.) Benth. 

Elymus canaliculatus (Nevski) 

Tzvel. 
E. nutans Griseb. 
Enkianthus deflexus (Griff.) 

Schneid. * 
Ephedra gerardiana var. 

sikkimensis Stapf * 



Epilobium conspersum Hausskn. 
E. leiospermum Hausskn. 
E. cf. wallichianum Hausskn. * 
E. williamsii Rawen 
Erigeron bellidioides 

(Buch.-Ham. ex D. Don) 

Benth. ex C.B.Clarke* 
E. kumaunensis (Vierh.) 

Wendelbo 
E. multiradiatus (Lindl. ex DC.) 

C. B. Clarke * 
Eriophy ton wallichii Benth. * 

Erysimum hieraciifolium L. 
Euonymus amygdalifolius 

Franch. 
£. /ng/du5 Wall. 
E. hamiltonianusWaW. * 
E. pendulus Wall. 
E. porphyreus Loesener + 



T caesp 


V 


W.204 


Tsem 


V,VII 


Z.807 


a f Ch herb pulv/a 


IX, X 


BL.2075,M.1102,Sm.s.n., 


Hros 




S.7158,W.175,184 


a f Ch herb pulv 


VI, VII 


D.445 


a H ros/a f Ch herb 


VII, IX 


P.G.206 


pulv 






a H ros/a f Ch herb 


X 


BL.2190,M.1098,Z.569 


pulv/G rhiz 






a Hros 


III,V 


By.365, D.333, Z.1694 


aHrept/afChherb 


IX, X 


BL.2119 


pulv 






a H scap/G rad 


V,VI 


By.583,590,591,dH.2962, 
M.880,P.s.n.,W.330, 
Z.s.n. 


a/e Mes/Mi P 


I 


P.s.n.,Sch.2442A,S.6574, 


scap/caesp 




Z.1850 


T scap/caesp 


II, V 


Z.1778 


a H scap 


V 


Z.683 


Tscap 


II, V 


By.205, 675, D.339, 
L.0180, M.889 


Tscap 


II, V 


Z.643 


a NP caesp/Ch suff 


II 


By.509, 611, 882, 1123, 
M.822,P.s.n.,Z.1654, 
1828 


Tscap 


II, III 


M.889a,P.s.n.,W.302, 
Z.1726 


a H caesp 


V,VI 


M.851a 


a H caesp 


VI 


P.s.n. 


s/a Mi P caesp 


I 


BL.2013, Sch.2336, Z.426 


pChfrut 


II, V, VI, IX 


By.736, BL.2069, D.439, 
M.993, P.R.333, G.293, 
Sch.2102, W.258, 
Sm.s.n. 


a H scap 


V,VI 


By.310, P.R.336 


a H scap 


7 


W.384, 392 


a H scap 


IV, VIII 


M.844 




? 


W.300 


aHsem 


VL 


D.325,Z.447,451,727 


aHsem 


II, V 


Z.1864 


aHsem 


V,VI 


By.294,M.876,Z.451, 
1565, 1622 


a G rhiz/scap 


X 


By.ll73,M.1007,W.236, 
Z.1510, Sm.s.n. 


Tscap 


II, V 


Z.450 


s NP caesp 


I 


S.6573 


s NP caesp 


I 


P.s.n. 


a Mi P scap 


I 


By.988 


s Mes P scap 


I 


S.6572 


a Mi P scap 


I 


Mc.43, S.5949 



244 


G. MIEHE 






E. tingens Wall. * 


s Mi P scap 


I, III 


S.4607 


Euphorbia himalayensis Klotzsch 


a G rad/scap 


II, V 


Sch.2086, 2291, 2324 


E. longifolia D. Don 


a G rad/scap 


II, V 


Sch.2069 


E. pseudosikkimensis (Hurusawa 


a G rad/scap 


II, V 


dH.2959 


& Ya. Tanaka) Radcliffe-Smith 








E, sikkimensis Boiss. 


a G rad/scap 


II, V 


Sch.2060 


E. stracheyi Boiss. 


a G rad/rept 


V, VI, IX 


B.5698,BL.2131,M.954, 
Z.490, 410, 537, Sm.s.n. 


E. wallichii Hook.f. * 


a G rad/scap 


II, V 


By.l01,1127,M.826, Z.37 


Euphrasia himalayica'Wettst . * 


Tscap 


VIII, IX 


By.415, Sm.s.n. 


E. platyphylla Pennell 


Tscap 


VII 


S.656 


E. secundiflora Pennell 


Tscap 


V 


P.s.n. 


Eurya acuminata DC. * 


s Mes P scap 


I 


S.4482 


Fagopyrum dibotrys (D. Don) 


aHsem 


II 


Z.766, 1862 


Hara 








F. esculentum Moench 


a H scap 


V 


By.304, 701 


Festuca ovina L. 


a H caesp 


VI 


Mc.289,W.183,242, 
Sm.s.n. 


F. polycolea Stapf 


a H caesp 


VI, VII 


D.373, S.550 


Festuca tibetica (Stapf) Alexeev + 


a H caesp 


X 


M.1072 


Ficus sarmentosa Buck. -Ham. ex 

Sm. 
Fragaria nubicola Lindl. ex 


s r PL frut 


I, IV 


f.M. 


a H caesp 


V 


By. 1129 


Lacaita * 








Fritillaria cirrhosa D. Don 


a G bulb 


III, V, VII 


Z.690, Sm.s.n. 


Galearis stracheyi (Hook.f.) P. F. 


aGrhiz 


VI, VII 


By. 131, D. 427 


Hunt* 








Galinsoga parviflora Cav. 


Tscap 


V 


Z.1765 


Galium aparine L. var. aparine 




III,V 


By.636, P.s.n. 


G. asperifolium Wall. 


Trept 


III 


W.426 


G. asperifolium var. sikkimense 


Trept 


IV 


P.R.275 


(Gandoger) Cufodontis 








G. asperuloides subsp. 


Trept 


III,V 


W.306 


hoffmeisterii (Klotzsch) Hara 








GaultheriafragrantissimaWdW. * 


s NP caesp 


III 


Sch.2050, Z.444, 1856 


G./iooA;en C.B.Clarke 


s NP caesp 


III 


BL.2022, S.4612 


G. nummularioides D. Don 


s Ch frut rept 


I 


M.820 


G. pyroloides Hook.f. & Thorns. 


s Ch frut rept 


III 


S.4616, Z.792 


ex Miq. * 








G. trichophylla Royle * 


s Ch frut rept 


III 


By.l09,M.970,S.4541, 
Z.623, 1785 


Gentiana algida Pall. * 


e H caesp/rept 


VIII 


S.6591 


G. algida var. nubigena (Edgew.) 


e H rept/sem 


VIII 


M.1075,P.G.203, 


Kusn. 






Sch.2313, Sm.s.n. 


G. algida var. parviflora (C. B. 


e H rept 


VIII 


Sm.s.n. , 


Clarke) Kusn. 






1 


G. algida subsp. przewalskii 


e H rept/sem 


VIII 


H.49 


(Maxim.) Kusn. 








G. argentea (D. Don) C. B. 


eHscap 


II, III 


E. Onyon 5 


Clarke 








G. capitata Buch.-Ham. ex D. 


eHros 


III 


Sw.525 


Don* 








G. crassuloides Bur. & Franch. 


e H caesp 


7 


Sm.s.n. 


G. depressa D. Don * 


e f Ch herb pulv 


V 


By.362,H.56,M.1117, 
P.147, 149,Sch.2316, 
Tilman s.n. 



PLANTS SOUTH OF MT EVEREST 



245 



G.e/we^// C.B.Clarke 
G.m/e/u: C.B.Clarke 
G. ludlowii Marq. 
G. micans C. B. Clarke 
G. ornata (G. Don) Griseb. 



G. ornata forma alba 

G. pedicellata (D. Don) Griseb. * 

G. phyllocalyx C. B. Clarke * 

G. prolata Balf.f. 

G. sikkimensis C. B. Clarke 

G. speciosa (Wall.) Marq. 

G. squarrosa Ledeb. + 

G. 5te//flto Turrill + 

G. strobilacea H. Sm. 

G. tubiflora (G. Don) Griseb. * 

G. Mmw/fl H. Sm. * 

G. vernayi Marq. 

Gentiana sp. 

Gentianella falcata (Turcz. ex 

Kar. & Kir.) H. Sm. 
G. glanduligera Airy Shaw 
G. pedunculata (D. Don) H. Sm. 
G. stellariifolia (Franch. ex 

Hemsl.) H. Sm. 
Geranium donianum Sweet * 

G. nakaoanum Hara * 

G. polyanthes Edgew. «fe Hook.f. ' 

G. refractum Edgew. & Hook.f. * 



Gerbera maxima (D. Don) 
Beauverd var. maxima 

G. nivea (DC.) Sch. Bip. 

Girardinia diversifolia (Link) 
Friis * 

Goodyera fusca (Lindl.) Hook.f. 

G. repens (L.) R. Br. * 

Gueldenstaedtia himalaica Baker 

Gymnadenia orchidis Lindl. 
Gypsophila cerastioides D. Don * 



Habenaria arietina Hook.f. 
Hackelia uncinata (Royle ex 

Benth.) C. E. C. Fischer 
Halenia elliptica D. Don * 
Hedera nepalensis K. Koch 
Hedychium densiflorum Wall. 
H. spicatum Smith * 
Hedysarum sikkimense Benth. ex 

Baker * 



aHsem 


VII 


Beer 25368 


a H caesp 


VIII 


BLM9548 


e H sem/caesp 


V, VI, VIII 


H.2,25,M.940,P.R.291 


e H sem/caesp 


V 


dH.2963, Sm.s.n. 


e H caesp/sem/e f 


V,VI 


By.597,H.35,M872, 


Ch herb pulv 




P.R.290,G.212,S.1673, 
6587, Sch.2314, Sharma 
E.465, W.220, Sm.s.n. 


e f Ch herb pulv 


VI 


M.1082 


H caesp 


V 


Sw.428 


a H sem/G rhiz 


V, VI, IX 


S.589 


e H caesp/rept 


V 


Sch. 2184 


a H scap 


? 


W.158 


a H rept 


i,ii 


P.B.9, 153,154,Sch.2081 




VI, IX 


D.2996, P.s.n. 


a H sem/caesp 


VI 


P.G.287,S.1674,1694, 
W.239 


eHros 


VII 


Sw.314 


eHros 


IX, X 


BLM9549 


e H ros/G rhiz 


X 


M. 1080, Sch.2314, S.6595, 
Sw.21-23, Sm.s.n. 


T caesp 


V, VII, IX 


H.24,M.1088,W.205 


aHros 


X 


M.1088 


T ros/sem 


VI, IX 


M.999, P.R.334, G.253, 
G.302, Sm.s.n. 


a H ros/sem 


? 


W.427 


T sem/scap 


IX 


M.lllO, P.s.n., W.221 


T caesp 


VI, VII 


S.1677 


aHsem 


VII 


By.l4, 152,D.335,401, 
M.947, S.7150, Sm.s.n. 


aHros 


VII 


By.ll46,D. 360, Sm.s.n. 


aHsem 


V 


By.275, P.G.329 


aHsem 


VII 


By.l92,234,463,BL.2096, 
D.420, H.37, Mc.347, 
Sm.s.n. 


a H sem/G rad 


II 


Z.1872 


a G rad/H ros 


V 


By.670, 787, P.s.n., W.353 


a H scap 


II 


Z.1874 


a G rad/rhiz 


III 


M.925,W.316 


a G rhiz 


III 


By.642 


a G rad/H caesp 


V-VII 


By.268,BL.2065,D.345, 
H.32,Mc.321, Sm.s.n. 


aGrad 


V-VII 


L.0193 


a H rept/a f Ch herb 


VIII 


By.316,BL.2165,M.979, 


pulv 




Mc.344 


a G bulb 


II, III, V 


H.70 


a H scap 


III,V 


BL.274 


Tsem 


V 


By.239,431,M.918,W.294 


s r PL f rut 


I, III 


Z.1838, 1840 


a G bulb 


I, II 


Sch.2042 


a G bulb 


I, II 


Sch.2043 


a H rept 


V-VII 


BL.2079,By.43,438, 
M.1024,Mc.310,S.7101, 
Sm.s.n. 



246 



G. MIEHE 



Helictotrichon sp. 


a H caesp 


V 


M.892 


Hemiphragma heterophyllum 


a H rept 


m 


dH.2978,Z.1819 


Wall. * 








Heradeum brunonis (DC.) C. B. 


aGrad 


? 


By.322,345 


Clarke 








H. candicans Wall, ex DC. 


aGrad 


I, III, V 


BL.2150, By.850, S.6598 


H. nepalense D. Don 


aGrad 


III, V 


M.883b 


H. wallichii DC. 


aGrad 


III,V 


By.809, M.883C 


Herminium josephii Reichenb.f . 


a G bulb 


VI, VII 


D.393 


H. lanceum (Thunb.) Vuijk * 


a G bulb 


V 


H.65a 


H. macrophyllum (D. Don) 


a G bulb 


V 


BL.2174,M.945,Mc.311 


Dandy 








H. pugioniforme Lindl. ex Hook.f. 


a G bulb 


V 


Mc.355 


Hierochloe flexuosa Hook.f. 


a H caesp 


? 


By. 520 


Hippophae tibetana Schlecht. * 


a/e Ch frut/pulv 


VIII 


By.428,dH.2989,f.M., 
P.G.305, f.Sch.(2268), 
2179 


Holboellia latifolia Wall. 


a el PL frut 


I, II 


S.4490 


H. latifolia var. angustifolia 


a el PL frut 


I, II 


Z.432, 473 


(Wall.) Hook.f. & Thorns. * 








Hydrangea heteromalla D. Don 


a Ch frut/r PL frut 


I 


Mc.269 


Hypecoum leptocarpum Hook.f. 


a H caesp/rept 


VII, VIII 


By.768, Sm.s.n. 


& Thorns. 








Hypericum choisianum Wall, ex 


a H caesp 


n 


Mc.399, S.7168 


N. Robson * 








H. himalaicum N. Robson 


a H scap 


n 


Mc.404 


Ilex dipyrena ^a\l. * 


s Mi P scap 


I 


Z.1675 


I. intricata Hook.f. 


s NP caesp 


m 


P.73, S.4611 


/. odorata Buch.-Ham. ex D. Don 


s Mi P scap 


m 


S.4610 


/. sikkimensis Kurz 


a Mi P scap 


I 


BL.2019 


Impatiens falcifer Hook . f . 


a H caesp 


? 


By.246, 376 


/. glandulifera Royle * 


Tscap 


II, V 


By.665 


/. puberula DC. 


a H scap 


I, IV 


D.328 


/. pulchra Hook.f. & Thorns. 


a H scap 


I, IV 


L.067 


/. sulcata Wall. * 


a H scap 


I, IV 


By.300, 573-575, W.341 


/. urticifolia Wall. * 


a H scap 


V 


f.M. 


Indigofera bracteata Grab, ex 


a H caesp 


II 


D.326 


Baker 








/. heterantha Wall, ex Brandis * 


a NP caesp 


n 


S.4622 


Inula cappa (Buch.-Ham. ex D. 


a/e Ch suff 


n 


Z.1881 


Don) DC. * 








/./looiten C.B.Clarke* 


a H scap 


n 


dH.2916,P.s.n.,S.6570, 
Z.1693, 1845 A 


/. nervosa Wall, ex DC. 


a/e G rhiz/H scap 


II 


dH.2919 


Iris kemaonensis D. Don ex 


a G rhiz 


V 


By.81 


Royle * 








Jasminum humile L. * 


s d PL frut 


n 


dH.2913, Z.666 


/. humile forma pubigerum (D. 


s d PL frut 


n 


By.860 


Don) Grohmann 








J. humile var. revolutum (Sims) 


s d PL frut 


II, V 


P.G.203 


Stokes 








Juncus benghalensis Kunth 


a G rhiz 


IV, VIII 


P.R.327 


J. clarkei Buchenau + 


a G rhiz 


7 


W.322 


/. concinnus D. Don 


a G rhiz 


IV 


M.868, P.B.55, 171 


J. grisebachii Buchenau 


a G rhiz 


? 


W.376 


/. himalensis Klotzsch * 


a G rhiz 


VIII 


M. 1029a, S.567 


/. leschenaultii Gay 


a H caesp 


IV 


Z.1724 



J. leucanthus Royle ex D. Don ' 

J. leucomelas Royle ex D. Don 
J. pseudocastaneus (Lingelsh.) 

Samuelsson 
/. sphacelatus Decaisne 
/. thomsonii Buchenau 



Juniperus indica Bertol. 



J. recurva Buch.-Ham. ex D. Don 

* 

J. squamata Buch.-Ham. ex D. 
Don* 

Kobresia caricina Willd. 

K. curvata Kiikenthal 

K. deasyi C.B. Clarke 

K. fissiglumis C. B. Clarke + 

K. macrantha Boeck. 

K. nepalensis (Nees) Kukenthal 

K. pygmaea (C. B. Clarke) C. B. 

Clarke 
K. schoenoides (C. A. Meyer) 

Steudel 
K. seticulmis Boeck. 

K. trinervis Boeck. 

Koenigia delicatula (Meisn.) Hara 

K. islandica L. 

K. nepalensis D. Don 

Lactuca lessertiana (DC.) C. B. 

Clarke * 
Lagotis kunawurensis (Royle ex 

Benth.) Rupr. * 
Lamium nepalense Hedge 
Lancea tibetica Hook.f. & 

Thorns. * 
Lasiocaryum densiflorum 

(Duthie) J. M. Johnston 
L. diffusum (A. Brand) J. M. 

Johnston 
L. munroi (C. B. Clarke) J. M. 

Johnston 
Leibnitzia nepalensis (Kunze) 

Kitam. 
L. ruficoma (Franch.) Kitam. 
Leontopodium brachyactis Gand. 

L. jacotianum Beauverd * 

L. monocephalum Edgew. 



PLANTS SOUTH OF MT EVEREST 

a G rhiz IV 



a H caesp 
a G rhiz 

a G rhiz 
a G rhiz 



VIII 
? 

VIII, X 
III, IV 



b N/Mi P scap/Ch III, V, VI 
frut 



247 

By.285, D.409, M.956, 
W.188a,252,Sm.s.n. 
D.409 
By.546 

D.377,M.1093,P.R.326 
By.222, P.G.264, S.7096, 

W.188, 190, Z.665, 1524, 

1540 
By.ll34,D.459,P.G.300, 

318,8.4513,4515,4528, 

7116 



b N/Mi P scap 


III,V 


By.80,P.3, 4,8.4529, 451^ 
W.410 


b Ch frut/rept 


VI 


By.ll68,Z.1699 


a H caesp 


VIII, X 


BL.2127 


a H caesp 


IV 


M.867,P.132 




VIII, X 


P.G.267 


a H caesp 


? 


W.269 


a H caesp 


VIII, X 


BL.2188 


a H caesp 


VI, VIII, IX, 
X 


M.s.n., 959, 988, 1071a 


a H caesp 


IX 


M.957, 1071b, P.R.332, 

W.237 


a H caesp 


VIII, X 


M.1053, 1104, nil 


a H caesp 


IV, VI, VII, X 


M.866, 1087, Mc.2032, 
P.133,134,R.226 


a H caesp 


VI 


BL.2197 


Trept 


IV 


M.916,P.s.n.,W.382 


Trept 


VIII 


W. 245, Z. 651 


Trept 


VIII 


M.981,Mc.312,W.381 


a H sem/G rad 


III, VI, VII 


M.982, 1023,W.251, 
Z.1529,8m.s.n. 


aHros 


VIII, X 


BL.2105,By.392,D.407, 
M.1069,8m.s.n. 


a H scap 


II 


L.0139 


aHros 


VI 


BL.2117,By.l3,D.442, 
8m.s.n. 


T caesp 


VI, IX 


By. 668 


T caesp 


VI, IX 


W.208 


aHsem 


7 


Beer 25543 


aHros 


V 


Z.403 


a H ros/G rad 


V 


dH.2945,Z.1754 


a H sem/f Ch herb 


V,VI 


M.881,942,W.403 


pulv 






G rhiz/ros/sem 


VI 


D.356, Z.1607, 1697, 
8m.s.n. 


a f Ch herb pulv 


IX 


BL.2139,D.428,H.48, 
M.1048, 1068, 1092, 
P.G.207, G.257, 8.6582. 
W.166,Z.1591,Sm.s.n. 



248 

L. nanum (Hook.f. & Thorns, ex 
C. B. Clarke) Hand.-Mazz. 

L. stracheyi (Hook.f.) C. B. 
Clarke ex Hemsley 

Lepidium capitatum Hook.f. & 

Thorns. 
Leptodermis kumanoensis Parker 

L. stapfiana H. Winkler 
Leucosceptrum canum Sm. * 
Leycesteriaformosa Wall. * 
Lignariella hobsonii (Pearson) 

Baehni * 
L. hobsonii subsp. serpens (W. 

W. Smith) Hara 
Ligularia amplexicautis DC. * 
L. fischerii (Ledeb.) Turcz. * 
Lilium nanum Klotzsch * 

L. nepalense D. Don * 
hinder a heterophylla Meisn. 
L. neesiana (Wall, ex Nees) Kurz 
L. pulcherrima (Nees) Benth. ex 

Hook.f. * 
Listera nepalensis Balakrishnan 
Litsea cubeba (Lour.) Pers. 
L. sericea (Wall, ex Nees) 

Hook.f. 
Lloydiaflavonutans Hara 
L. longiscapa Hook * 
L. serotina (L.) Reichenb. * 
L. serotina var. parva (Marquand 

& Shaw) Hara 
Lobelia seguinii H. Lev. & Van. 
Lomatogonium brachyantherum 

(C. B.Clarke) Fernald 
L. carinthiacum (Wulf.) 

Reichenb. * 
L. chumbicum (Burkhill) H. Sm. 
L. graciliflorum H. Sm. 
L. sikkimense (Burkhill) H. Sm. 
Lonicera acuminata Wall. 
L. angustifolia Wall, ex DC. 

L. cyanocarpa var. porphyrantha 

Marquand & Shaw * 
L. /iwpidaPall. exWilld. * 
L. hispida var. setosa Hook.f. & 

Thorns. 
L. lanceolata Wall. 

L. litangensis Batahn 

L. myrtillus Hook.f. 8c Thoms. * 

L. obovata Royle ex Hook.f. & 

Thoms. * 
L. rupicola Hook.f. & Thoms. * 

Luzula sp. 



G. MIEHE 






a f Ch herb pulv 


IX 


BL.2125,M.1041 


a H caesp 


n 


By.389, dH.2953, M.833, 
P.lll,112,Sch.2176, 
Z. 1651, 1821 


Trept 


DC 


M.1116 


a NP caesp 


m 


S.7160 


a NP caesp 


II 


S.4618 


a NP caesp 


II 


Z.1858 


a NP caesp 


II 


P.96, Z.743 


Trept 


VIII 


W.331 


Trept 


V 


BL.2207 


aHsem 


III,V 


By. 171 


a H scap 


II, V 


By.286, L.0182 


a G bulb 


II, V 


BL.2056, 2074, By. 150, 
S.4775 


a G bulb 


I,II,V 


By.l76,179,Sch.2248 


s Mes P scap 


I 


S.6566 


a/s Mes P scap 


I 


S.6571,6599 


a/s Mes P scap 


I 


Z.260 


aGrad 


? 


W.339 


a Mes P scap 


i 


S.6567 


s Mes P scap 


I 


S.5942 


a G bulb 


V 


Mc.279 


a G bulb 


V 


Beer 25375, Sm.s.n. 


a G bulb 


VI, VII 


BL.2962, By.308, D.375 


a G bulb 


VI 


BL.2170, M.944, Mc.326 


a H scap 


II 


Z.1863 


a H caesp 


VIII, IX 


H.34, P.R.347 


aHsem 


VI-IX 


By.ll74,M.1056,P.G.223, 
R.329, S.6593, W.250 


aHsem 


VI 


M.946, W.402 


aHsem 


? 


W.292, 400 


aHsem 


V 


W.420 


a Ch frut 


n 


Mc.393 


a Mi P caesp 


III 


BL.2083, 2152, By.68, 873, 
M.935, Z.429 


a Ch frut 


VII 


Mc.305 


a NP caesp 


VI, VII 


D.434, Sm.s.n. 


aChfrut 


VI, VII, IX 


BL.2104,M.1050,Mc.350 


a NP caesp 


m 


BL.2023,By.51,282,557, 
1002, D.452, M.934 


aChfrut 


VI, VII 


Z.520 


aChfrut 


VI, VII 


By.425, D.418 


a Ch frut 


V-VII 


BL.2185, By.903, P.R.357, 
Sch.2078, Z.614, Sm.s.n. 


a Ch/NP caesp 


III,V 


By.37, M.985, P.R.358, 
Z.509, 1530, Sm.s.n. 


H caesp 


m 


P.R.341 



PLANTS SOUTH OF MT EVEREST 



249 



Lyonia ovalifolia (Wall.) Drude 



Lysimachia prolifera Klatt 

Malaxis cylindrostachya (Lindl.) 

Kuntze 
M. muscifera (Lindl.) Kuntze 
Mandragora caulescens subsp. 

flavida Grierson & Long * 
Meconopsis bella Prain * 
M. horridula Hook.f. «fe Thorns. * a H sem 



M. paniculata Prain * 
M. simplicifolia (D. Don) Walp * 
M. sinuata Prain * 
Megacodon stylophorus (C. B. 

Clarke) H. Sm. * 
Meliosma dilleniifolia (Wall, ex 

Wight & Am.) Walp. 
Michelia doltsopa Buch.-Ham. ex 

DC* 
Microcaryum pygmaeum (C. B. 

Clarke) J. M. Johnston 
Micromeria biflora (Buch.-Ham. 

ex D. Don) Benth. * 
Microsisymbrium axillare 

(Hook.f. & Thorns.) O.E. 

Schulz 
Microula pustulosa (C. B. 

Clarke) Duthie 
M. sikkimensis (C. B. Clarke) 

Hemsl. * 
Miscanthus nepalensis (Trin.) 

Hackel 
Myricaria davurica (Willd.) 

Ehrenb. 
M. rosea W. W. Sm. * 



Nannoglottis hookeri (C. B. 
Clarke ex Hook.f.) Kitam. * 



Nardostachys grandiflora DC. * 
Neilla rubiflora D. Don 
Neolitsea pallens (D. Don) 

Momiyama & Hara ex Hara * 
Neottianthe calcicola (W. W. 

Smith) Schlechter 
N. secundiflora (Hook.f.) 

Schlechter 
Nepeta coerulescens Maxim. 
N. lamiopsis Benth. ex Hook.f. 



Notholirion macrophyllum (D. 
Don) Boiss. * 



a/s N/Mi P scap 


III 


f.M. 


a/s N/Mi P scap 


III 


M.858,P.G.358,Z.1852 


a H caesp/rept 


I, III 


Z.781 


a G bulb 


II, V 


Mc.397 


a G bulb 


V 


By.332, H.15 


a G rad/ros 


V 


Z.688 


a H scap 


III,V 


Beer 25425, BL.2161 


aHsem 


V-VII, IX 


By.451,H.51,P.s.n., 
Sch.2177,2351,W.280, 
Sm.s.n. 


aHsem 


III,V 


S.649, W.395 


a H scap 


III, VII 


By.28, Mc.348, Z.594 


a H scap 


VII, VIII 


Beer 25445 


a H scap 


III,V 


BL.2153,By.271,L.0183, 
Z.1703 


a Mi P scap 


I 


S.4620 


s Mes P scap 


I 


S.4478 


Tros 


V, VI, IX 


M.s.n. 


a H caesp 


II 


Z.400 


aHsem 


II, V 


Z.258 


a H caesp/scap 


V,VI 


BL.2156 


a H scap 


V 


BL.2183, D.367, M.887, 
Sm.s.n. 


a H caesp 


II 


Sch.2063,Z.1859 


sChfrut 


VIII 


Z.510 


s Ch frut rept 


VIII 


By.350,dH.2971,M.969, 
P.s.n., Sm.s.n., W.432 


a H scap/G rad 


III 


By. 220, Einarsson 9, 
L.0178, M.870, Mc.444, 
P.s.n., S.3820, 7130, 
Z.635,687 


aHsem 


V,VI 


L.0212 


a NP caesp 


II 


BL.2081 


s Mi P scap 


I, II 


S.4482 


a G bulb 


II, V, VII 


H.26,M.963,P.16 


a G bulb 


II, V 


H.61,P.16 


a H scap 


V 


B.5778 


a H rept/scap 


V 


B.5636,By.52,D.385, 
H.16,L.0170,M.905, 
Mc.359, P.s.n., Z.1766 


aGrad 


III, V-VII 


By.217,D.331,Sch.2105 



250 

Omphalodes trichocarpa 

Maxim. + 
Orchis diantha Schlechter + 
Oreorchis foliosa (Lindl.) Lindl. * 
O. micrantha Lindl. 
Oreosolen wattii Hook.f. * 

Oxalis corniculata L. * 

O. corniculata var. villosa (Bieb.) 

Hohen + 
Oxyria digyna (L.) Hill * 

Oxytropis arenae-ripariae Vass. 
O. nepalensis Vass. 
O. williamsii Vass. * 

Paris polyphylla Smith * 
Parnassia chinensis Franch. 
P. kumaonica Nekrassova 
P. nubicola Wall, ex Royle * 
P. pusilla Wall, ex Arn. 

Parochetus communis 
Buch.-Ham ex D. Don * 

Pedicularis bifida (Buch.-Ham. 
ex D. Don) Pennell 

P. brevifolia D. Don 

P. clarkei Hook.f. 

P. confertiflora Prain 

P. elwesii Hook.f. 

P. flexuosa Hook.f. 
P. furfuracea Wall, ex Benth. 
P. gracilis Wall, ex Benth. 
P. longiflora var. tubiformis 

(Klotzsch) Tsoong * 
P. megalantha D. Don 
P. microcalyx Hook.f. 
P. mollis Wall, ex Benth. 
P. muscoides L. 
P. nana C. E. C. Fischer 
P. oederi suhsp. oederivar. 

heteroglossa Prain * 
P. pseudoregeliana Tsoong 
P. punctata Deer. + 
P. rhinanthoides Schrenk. 
P. roy lei Mdixim. 
P. Scully ana Prain ex Maxim. * 

P. siphonanta D. Don 
P. trichoglossa Hook.f. * 

P. wallichii Bunge 
Pegaeophyton minutum Hara 

P. scapiflorum (Hook.f. & 
Thoms.) Marq. & Shaw * 

Pentapanax leschenaultii (DC.) 
Seem. * 



G. MIEHE 








? 


By.41,213,374,892 


a G rhiz 


V,VI 


Mc.294, BL.2048 


a G bulb 


I 


BL.2026, Mc.292 


a G bulb 


1,11 


BL.2025 


aHros 


IX 


BL.2124,M.1095,W.230, 
Z.535, Sm.s.n. 


a H rept 


I 


Z.1842 


a H rept 


? 


Onyon 9 


aGrad 


VIII 


By.287,M.1004,P.s.n., 
S.7598, Z.599, Sm.s.n. 


a H caesp 


VIII 


S.7109 


a H caesp/rept 


VI 


BL.2107 


a H caesp 


IX 


By.758 


a G rhiz 


I 


Z.449 


aHros 


IV 


M.s.n., Sm.s.n. 


aHros 


IV, VIII 


By.346, M.864, Mc.328 


aHsem 


IV, VIII 


By.518, L.0128, M.900 


aHros 


IV, VIII 


BL.2184,H.ll,Mc.352, 
Sm.s.n. 


Trept 


V 


By.s.n.,W.423 


a H caesp/scap 


? 


By.361 


aHsem 


VI 


P.R.294 


a G rad/H scap 


V 


B.5649 


Tsem 


VIII 


H.29 


H caesp/G rad 


VII, VIII 


BL.2093,By.258,D.351, 
W.286 


aHsem 


m 


Beer 25339 


aHsem 


ra 


BL.2030, Mc.268 


a H scap 


V 


B.5647,By.371,M.896 


a H ros/G rad 


VIII 


H.31,BL.2181,By.277 


a H scap/G rad 


III, V, VII 


By.289,641,M.846 


G rad/scap 


VII 


BL.2137 


a H caesp/scap 


II, V 


B.5697,By.247,H.l,P.55 


a H ros/G rad 


VIII 


BL.2129, Mc.366 


a H ros/sem 


VI, VII 


D.433 


G rad/H sem 


VIII 


D.411 


aHros 


II, VII 


BL.2086,2201,S.4603 


a H caesp/scap 


? 


By.460 


a H caesp 


7 


By. 122 


a H caesp 


VIII 


By.440, D.382 


a H scap 


VII 


By.422,Shresta5661, 
Sm.s.n. 


a H ros/caesp/G rad 


VIII 


BL.2180, By.434, D.381 


a H scap 


VII 


By.368, D.350, L.0185, 
M.965, P.G.234 


aHsem 


vn 


Mc.285 


a H caesp/a f Ch 


VIII 


BL.2088, Mc.388 


herb pulv 






aGrhiz/afChherb 


IX, X 


M.1099,Mc.335,373 


pulv 






a Mi P caesp 


I 


BL.2158 ^..- 



Peracarpa carnosa (Wall.) 

Hook.f. & Thorns. 
Peristylus elisabethae (Duthie) 

Gupta 
Peristylus fallax Lindl. 
Persicaria glacialis (Meisn.) Hara 
P. microcephala var. 

sphaerocephala (Wall, ex 

Meisn.) Hara 
P. polystachya (Wall, ex Meisn.) 

H. Gross 
P. runcinata (Buch.-Ham ex D. 

Don) H. Gross 
Philadelphus tomentosus Wall, ex 

G. Don * 
Phleum alpinum L. 
Phlomis macrophylla Wall, ex 

Benth. 
P. rotata Benth. ex Hook.f. * 



Phyllanthus clarkei Hook.f. 

Physospermopsis obtusiuscula 

(DC.) C. Norman 
Picrorhiza scrophulariiflora 

Pennell 
Pieris formosa (Wall.) D. Don * 

Pilea racemosa (Royle) Tuyama 
P. symmeria Wedd. 
Pinguicola alpina L. * 
Pinus wallichiana A. B. Jackson * 
Piptanthus nepalensis (Hook.) D. 

Don* 
Platanthera clavigera Lindl. 
P. latilabris Lindl. 
Pleione hookeriana (Lindl.) J. 

Moore 
Pleurospermum angelicoides 

(DC.) C. B. Clarke 
P. apiolens C. B. Clarke 

P. benthamii (DC.) C. B. Clark^'- 
P. brunonis (DC.) C. B. Clarke 
P. aff. corydalifolium Aitch. & 

Hemsl. + 
P. dentatum (DC.) C. B. Clarke 
P. hookeri C.B. Clarke 
P. rotundatum C. B. Clarke 
Poa annua L. 
P. hirtiglumis Hook.f. 
P. pagophila Bor 
P. polycolea Stapf 
Podophyllum hexandrum Royle * 
Polygonatum cf . cirrhifolium 

(Wall.) Royle * 
P. hookeri Baker * 



PLANTS SOUTH OF MT EVEREST 


231 


T rept/scap 


I 


Z.778 


a G bulb 


V 


H.60 


a G bulb 


n 


Mc.396 


Trept 


VIII 


M.917, W.244 


a H rept 


I, II 


Z.737 


a G rhiz 


I, II 


P.s.n.,Sch.2036,2039 




I, II 


P.s.n. 


a NP caesp 


I, II 


L.2089, S.4630, Z.733, 
1829 


a H caesp 


X 


P.G.311,W.428 


a H scap 


V 


By.585, P.R.310, Sch.2190, 
2280, Z.776 


aHros 


IX 


BL.2130,M.1055, 
P.G.242, Z.307, 578, 
Sm.s.n. 


a Ch frut/a NP 


II, V 


M.819 


caesp 






aGrad 


IX 


M.1036 


a G rhiz/ros 


VII 


BL.2051,Mc.280 


s NP scap 


I, III 


By.911,M.857, P.s.n., 
Z. 1719, 1937 


a G bulb 


III 


W.333 


a G rhiz 


I 


BL.2014 


aHros 


IV 


Beer 25369,8.660 


b Mes P scap 


II 


M.s.n. 


a Mi P scap 


II, III 


By.ll75, P.s.n., Sch.2082 


a G bulb 


V 


H.65 


a G bulb 


V 


H.66, L.0171 


fac a GE bulb 


I 


M.s.n. (Gottingen, Bot. 
Garden) 


aGrad 


V 


P.s.n. 


aGrad 


V,VII 


M.910a, 937, 987, 1016, 
P.s.n. 


aGrad 


III 


By.644, L.7, M.921 


aGrad 


II 


M.847 


aGrad 


? 


By.656 


aGrad 


? 


By.663, 831 


aGrad 


VIII 


By.449,M. 1085, Sm.s.n. 


aGrad 


III.V 


By.663, 831, M.910b 


T caesp 


VII, X 


Sm.s.n. 


a H caesp 


? 


W.243 


a H caesp 


X 


M.1103, Sm.s.n. 


a H caesp 


VI, IX, X 


P.G.306 


a G bulb 


V 


Z.618 


a G rhiz 


III,V 


S.7124 


a G rhiz 


VI, V 


BL.2066,By.880,M.941, 
Sm.s.n. 



252 

P. cf. kingianum Colett & Hemsl. a G rhiz 

P. verticillatum (L.) All. 

Ponerorchischusua(J).T>ovi)So6 a G bulb 



Potentilla argyrophylla Wall, ex 

Lehm. * 
P. argyrophylla Wall, ex Lehm. 

var. argyrophylla 
P. argyrophylla var. 

atrosanguinea (Lodd.) Hook.f. 

P. caliginosa Soj^k 

P. coriandrifolia D. Don * 

P. cuneata Wall, ex Lehm. * 



P. eriocarpa Wall, ex Lehm. * 
P. fruticosa var. rigida (Wall, ex 
Lehm.) Wolf* 



P. fulgens Wall, ex Hook. 

P. grifftthii Hook.f. 

P. cf . kleiniana Wight 

P. leschenaultiana Seringe 

P. leuconota D. Don 

P. microphylla D. Don * 

P. microphylla var. achilleifolia 

Hook.f. 
P. microphylla D. Don var. 

microphylla 
P. monanthes Wall, ex Lehm. 
P. peduncularis D. Don * 
P. polyphylla Wall, ex Lehm. 
P. saundersiana Royle 

Primula alpicola var. luna (Stapf) 

W. W. Sm. & Hetcher + 
P. atrodentata W. W. Sm. * 

P. buryana Balf.f. 

P. calderiana subsp. strumosa 

(Balf.f. & Cooper) A. J. 

Richards * 
P. capitata Hook.f. subsp. 

capitata 
P. capitata subsp. crispata (Balf.f. 

&W.W. Sm.)W.W. Sm. & 

Forrest 
P. caveana W. W. Smith * 



P. concinna Watt 
P. denticulata Sm. * 
P. deuteronana Craib 



G 


r, MIEHE 






a G rhiz 




I 


P.s.n. 


a G rhiz 




m 


By.878, M.926, Mc.303, 
Z.669 


a G bulb 




III,V 


By.226 


aHsem 




vn 


D.403, S.4774 


aHsem 




VI 


P.G.255, Z.586 


aHsem 




n,v 


f.M. 


aHsem 




DC 


Sw.258 


aHsem 




VI, IX 


D.363,Mc.383,P.35 


aHrept/fChherb 


II, V, VII 


By.l96,D.329,M.874, 


pulv 






P.33, 34, Sch.2035, 
W.439, Z.468, 613 


a G rhiz 




VII, VIII, X 


D.425,W.438,Sm.s.n. 


a Ch frut 




VII, X 


BL.2159,2533,H.21, 
D.329, M.852, 1079, 
P.G.294, G.724, S.4752, 
Sch.2033,2034,Z.613 


aHsem 




II, V, VI, IX 


M.821, 878, Z. 1659 


a H caesp 




V 


P.G.338 


aHsem 




V 


M.888 


aHsem 




V-VII 


By.942, D.348 


aHsem 




III,V 


BL.2053,M.974,W.418 


a f Ch herb pulv 


IX 


By.l097,Mc.251 


afChherb 


pulv 


IX 


D.349,M.1038,Z.619, 
665a 


afChherb 


pulv 


IX 


M.967, 1073 


aHsem 




VI 


BL.2108, Z.580a 


aHsem 




VI 


D.364, Z.656 


a H sem/G rhiz 


V,VI 


By.233, S.4689, 4787 


aHsem 




V 


By.45, 208, M. 1025, Z.500, 
628,685,1610a 


aHros 




vm 


Sch.2015 


aHros 




VllI 


By.417, S.4817, Z.269, 340, 
512, 563, Sm.s.n. 


aHros 




III, VIII 


BL.2054, Mc.378 


aHros 




V 


Beer 25354, Mc.281 


aHros 




V 


Sch.2012 


aHros 




vn 


Beer 25408 



aHros 



DC 



aHros 


VIII, IX 


aHros 


I,III,V 


aHros 


DC 



BL.2123A, P.G.283, 

S.6596,W.261,Z.1508, 

Sm.s.n. 
M. 1059, 1109, Z.1505, 

Sm.s.n. 
By.l29, P.s.n., Sch.2198, 

2292, Z.370, 402, 505 
S.4530, Z.815C 



PLANTS SOUTH OF MT EVEREST 



253 



P. dickieana Watt 


aHros 


V,VII 


Beer 25352 


P. gambeliana Watt 


aHros 


III,V 


By.390, BL.2037, Z.815 


P. geraniifolia Hook.f. * 


aHros 


I, III 


BL.2173, Mc.390, Z.726, 
1747 


P. glabra Klatt 


aHros 


V 


Z.814 


P. glomerata Pax. * 


aHros 


III,V 


By.352,649,dH.2983A, 
H.57,M.894,991,Z.816, 
Sm.s.n. 


P. gracilipes Craib 


aHros 


m 


S.4526 


P. hookeri Watt 


aHros 




Beer 25374 


P. listen King ex Hook.f. 


aHros 


I 


S.6975,Z.779 


P. macrophylla D. Don * 


aHros 


VII 


BL.2097, W.278 


P. macrophylla var. 


aHros 


VIII 


Z.1463 


moorcroftiana (Wall, ex Klatt) 








W. W. Sm. & Fletcher 








P. megalocarpa Hara 


aHros 


VIII 


Beer 25373, BL.2192, 
dH.2986, M.984, 
P.G.284,Z.1565, 
Sm.s.n. 


P. muscoides Hook.f. ex Watt 


a f Ch herb pulv 


VII 


Mc.389 


P. obliqua W. W. Sm. * 


aHros 


V, VIII 


Beer 25353, Mc.382 


P. primulina (Sprang.) Hara * 


aHros 


VI, VIII 


By.442,BL.2094,D.383, 
432, M.972, W.259, 
Z.s.n., 802, Sm.s.n. 


P. pulchra Watt 


aHros 


VIII 


BL.2039, S.7090 


P. reticulata Wall. 


aHros 


VIII 


Mc.406 


P. rotundifolia Wall. * 


aHros 


VII, VIII 


By.436,D.361,M.909, 
Mc.278, L.0187, Z.506, 
636, 1700, Sm.s.n. 


P. scapigera (Hook.f.) Craib 


aHros 


III 


Z.800 


P. sikkimensis Hook.f. * 


aHros 


VIII 


By.547, 592, 605, 636, 649, 
1500, D.369, Sm.s.n. 


P. stuartii Wall. * 


aHros 


vn 


Mc.386 


P. tanneri subsp. nepalensis (W. 


aHros 


m 


Z.799 


W.Sm.) A. J. Richards 








P. tenuiloba (Watt) Pax 


Tros 


VIII 


BL.2191 


P. uniflora Klatt 


aHros 


VIII, IX 


Beer 25360, S. 1649 


P. walshii Craib * 


aHros 


VI, IX 


P.s.n.,Z.282,511,522, 
Sm.s.n. 


P. wollastonii Balf .f . * 


aHros 


III, IV, VIII 


BL.2101,2136,By.274, 
D.402,H.18,M.842, 
P.R.312,Z.1626, 1807, 
Sm.s.n. 


Prunella vulgaris L. 


Tscap 


? 


By.585,W.414 


Prunus rufa Hook.f. * 


a Mi P scap 


m 


Sch.2256 


Pseudostellaria heterantha 


a G bulb 


? 


By.828 


(Maxim.) Pax 








P. heterantha var. himalaica Ohwi 


a G bulb 


VI, VII 


D.344 


Pueraria peduncularis (Benth.) 


a/e d PL suff 


II 


dH.2911A 


Grab. 








Pyrola sikkimensis Knsa 


aHsem 


I 


Z.483 


Quercus glauca Thunb. var. 


s Mes P scap 


I 


D.320 


glauca * 








Q. lanata Sm. * 


s Mes P scap 


I 


S.4539 


Q. semecarpifolia Sm. 


s Mes P scap 


I, II 


f.M. 



Rabdosia lophanthoides var. a H scap 

gerardiana (Benth.) Hara 



U 



Z.1879 



254 

Ranunculus adoxifolius 

Hand.-Mazz. * 
R. brotherusii Freyn var. 

brotherusii * 
R. brotherusii var. tanguticus 

(Maxim.) Tamura 
R. diffusus DC. 
R. hirtellus Royle ex D. Don * 
R. hyperboreus Rottb. 
R. oreionannos Marq. & Shaw 
R. pedatifidus Sm. + 
R. pegaeus Hand.-Mazz. 
R. pulchellus C. A. Meyer * 

R. pulchellus var. ellipticus 

Tamura + 
R. pulchellus var. sericeus 

Hook.f. & Thoms. 
R. pulchellus var. stracheyanus 

(Maxim.) Hand.-Mazz. 
Ranunculus sp. 
Rheum acuminatum Hook.f. & 

Thoms. ex Hook. 
Rhodiola amabilis (H. Ohba) H. 

Ohba 
R. bupleuroides (Wall, ex 

Hook.f. & Thoms.) Fu* 

R. crenulata (Hook.f. & Thoms.) 

H. Ohba 
R. cretinii (R.-Hamet) H. Ohba 

R. discolor (Franch.) Fu 
R. fastigiata (Hook.f. & Thoms.) 
Fu* 

R. himalensis (D. Don) Fu * 

R. humilis (Hook.f. & Thoms. 

emend. R.-Hamet) Fu 
R. ovatisepala (R.-Hamet) Fu 
R. prainii (R.-Hamet) H. Ohba 
R. quadrifida (Pallas) Fischer & 

Meyer * 
R. sinuata (Royle ex Edgew.) Fu 



R. wallichiana (Hook.) Fu 



Rhododendron anthopogon * D. 

Don var. anthopogon pulv 

R. anthopogon var. hypenanthum s NP caesp/s Ch frut VII 

(Balf.f.) Hara pulv 

R. arbor eum Smith * s Mes P scap I-III 



G. MIEHE 






aHros 


m 


S.7091 


aHros 


VII, VIII 


D.338,H.38,Sm.s.n. 


aHros 


VIII, IX 


W.202 


aHros 


I, II 


Z.745 


aHros 


VII, VIII 


W.458, Z.582, Sm.s.n. 


aHros 


VIII 


Sm.s.n. 


aHros 


VIII 


BL.2080, 2189, Sm.s.n. 


aHsem 


V 


P.R.284 


a H ros, fac hyd 


VIII 


BL.2146,W.197 


aHros 


VIII 


By.230,BL.2076,M.1113, 
1115,P.s.n. 


aHros 


ni 


S.7092 


aHros 


VIII 


Sm.s.n. 


aHros 


IX 


BL.2106,D.406,S.7110, 
Z.596 


aHros 


? 


W.305 


a H sem/G rad 


VII 


By.653,L.02O3,M.10O2, 
S.7134 


a Ch 1 succ 


? 


By.780 


a Ch 1 succ/G rad 


IV, IX 


Beer 25514, BL.2154, 
By.l80,M. 1017, 1044, 
P.G.233, Sm.s.n. 


a Ch 1 succ 


VIII, X 


By.779,M.1063,W.168, 
Sm.s.n. 


a Ch 1 succ/G 


VIII 


BL.2077 


rhiz/caesp 






a Ch 1 succ/caesp 


IX 


L.0211,Mc.349,S.4615 


aChlsucc/afCh 


IX, X 


BL.2084, 2085, 2095, 


herb pulv 




By.49,M.1105,Mc.339, 
Z.1514 


a Ch 1 succ 


DC 


By.408, D.378, Sch.2180, 
Z.701, Sm.s.n. 


a Ch 1 succ 


IV 


M.863 


e Ch 1 succ 


IV 


H.13,68,M.841 


a Ch 1 succ 


VII, VIII 


D.396, 397 


a Ch 1 succ 


X 


M.1062, 1096 


a Ch 1 succ 


III, IV 


By.702, 793, 794, M.838, 
Mc.270,P.103, 
Z.1628 


a Ch 1 succ 


IV, VIII 


H.27,58,M.923,P.R.301, 
s.n. 


s NP caesp/s Ch frut 


VII 


M.951 



R. arboreum var. cinnamomeum s Mes P scap 

(Wall, ex G. Don) Lindl. 
R. barbatum Wall, ex G. Don * s Mi P caesp 



I-III 
I, III 



Z.562, 1552, 1812, Sm.s.n. 

By.30,31,972,M.823, 

Sch.2297, Z.498 
Mc.46, Sch.2187, Z.271, 

371,497, 1784 
By.554, 605, 606, S.4525, 

Z.1848 



R. campanulatum var. wallichii 

(Hook.f.) Hook.f. * 
R. campy locarpum Hook.f. 



R. ciliatum Hook.f. * 
R. cinnabarinum Hook.f. 

R. dalhousiae Hook.f. * 
R. fulgens Hook.f. 

R. hodgsonii Hook.f. * 



R. lepidotum Wall, ex G. Don 



R. lepidotum var. eleagnoides 

(Hook.f.) Hook.f. 
R. lindleyi'T. Moore * 
R. nivale Hook.f. * 



R. setosum D. Don * 



R. triflorum Hook.f. * 

R. wightii Hook.f. * 

Ribes gtaciale Wall. 

R. grifftthiiHook.f. &Thoms. * 

R. himalense Royle ex Decne. 

R. luridum Hook.f. & Thorns. 

R. orientale Desf . * 

R. takare D.Don* 

R. takare forma desmocarpum 

Hara 
R. v///norwH Janczew. 
Rodgersia nepalensis T. A. Cope 

ex CuUen * 
Rosa macrophylla Lindl. * 

R. sericea Lindl. * 

Roscoea alpina Royle * 
Rubus biflorus Buch.-Ham. ex 

Smith 
R. fockeanus Kurz 
R. hypargyrus var. niveus (Wall. 

ex G. Don) Hara 
R. nepalensis (Hook.f.) Kuntze * 
R. paniculatus Smith * 
Rumex nepalensis Spreng. * 



Sabia campanulata Wall, ex 
Roxb. * 



PLANTS SOUTH OF MT EVEREST 

sMiPcaesp I, III 



s Mi P caesp 



III 



s NP caesp/s Ch frut I, III 
s Mi P caesp I 



255 
Mc.47, S.4527, Sch.2252 

By.l00a,s.n.,M.856,931, 
Mc.49, Sch.2188, S.2252, 
4602, Z.373, 412, 499 
Mc.57, S.4613, Z.793 
By.l86,BL.2031,Mc.277, 
S.4532 



s NP caesp/fac E 


I 


Z.463 


s NP caesp/s Mi P 


III 


M.930, Mc.288, S.4524 


caesp 






s Mi P caesp 


III 


By.685, 686, 968, Mc.48, 
S.4521,Sch.2343, 
W.411,Z.645 


s/a NP caesp 


I-III, V 


By.184,296,385,659, 
1008, 1109, D.347, 
M.853,860,Sch.2281, 
Z.673, 1669, 1782, 
1783B 


s/a NP caesp 


V,VII 


Sm.s.n. 


s NP caesp/fac E 


I 


Mc.39, S.4531 


s Ch frut/s Ch frut 


VII, IX 


By.915, D.430, M.996, 


pulv 




Sch.2269, Z.523A, 538, 
1484, 1811, Sm.s.n. 


s NP caesp/s Ch frut 


VII 


By.688, M.950, Sch.2048, 


pulv 




P.s.n.,W.442,Z.504, 
590, 1810, Sm.s.n. 


s NP caesp/s Ch frut 


I, II 


Mc.40, Sch.2048, S.4520, 
Z.431 


s Mi P caesp 


III 


Z.1784A 


a NP caesp 


III,V 


M.920 


a NP caesp 


ni 


S.4534 


a NP caesp 


I 


Z.414, 462 


a NP caesp 


VII 


D.355 


a NP caesp 


I 


Z.430 


a NP caesp 


ra 


S.5947 


a NP caesp 


m 


S.6982 


a NP caesp 


V 


Z.616 


eGrad 


n 


L.0152, S.4619 


a NP caesp 


III,V 


By.5, dH.2976, L.0168, 

P.37 


a NP caesp 


III,V 


By.l7,dH.2938,M.855, 
P.38, 39, R.278, 305 


aGrad 


I-III, V 


f.M.,By.l8,327,372 


a NP caesp 


7 


By.579 


a Ch frut rept 


V 


Z.692 


a NP caesp/a PL 


V 


dH.2990, M.904, P.40 


frut 






a Ch frut rept 


1, 11 


dH.2931 


a d PL frut 


I, IV 


dH.2931 


aGrad 


II, V, X 


By.215,395,P.s.n., 
Z.1720, Sm.s.n. 


a d PL frut 


I 


BL.2018,Z.481,489 



256 


G. MIEHE 






Salix calyculata Hook.f. ex 


a Ch frut rept 


VII 


B.5678, BL.2134, By.550, 


Anderss. * 






D.387,388,M.924,952, 
Sm.s.n. 


S. daltoniana Anderss. 


aChfrut/aNP 


III, VII 


By.6,446,D.457,M.1003, 




caesp 




W.343, Z.376 


5. disperma Roxb. ex D. Don 


a Mi P scap 


I 


Z.471 


S. eriostachya Wall, ex Anderss. 


a Ch frut rept 


VII 


By.l89,531,W.311 


S. hylematica Schneid. 


a Ch frut rept 


VII 


By.543,D.354,437, 
W.431,Z.1518 


5. cf. insignis Anderss. + 


a NP caesp 


IX 


P.R.344 


S. lindleyana Wall, ex Anderss. 


a Ch frut rept/f pulv 


III, VII 


D.436,M.962, 990, 1005, 
Sm.s.n. 


5. longifora Anderss. 


a Mi P scap 


I 


Z.387 


S. luctuosa H. L6v. 


a NP caesp 


i,v 


Z.388, 491 


5. serpyllum Anderss. 


a Ch frut rept 


VII 


Z.1518 


5. sikkimensis Anderss. * 


a NP caesp 


VII 


BL.2118,W.419 


S. tetrasperma Roxb. 


a NP caesp 


I 


Z.479 


Salix sp. 


a NP caesp 


? 


dH.2987 


5a/a sp. 


a Mi P scap 


in 


M.922 


Salvia campanulata Wall, ex 


aHsem 


V 


By.204,S.7141,Z.371A, 


Benth. * 






374, 407, 654, 1813 


5. castanea Diels 


aHsem 


V 


By.973,P.s.n.,Sch.2055, 
2057 


5. /wfl/M Royle ex Benth. * 


aHsem 


V 


BL.2151,By.l94,D.447, 

M.825 


Sanguisorba diandra (Hook.f.) 


a H scap 


III-V 


M.871,P.42 


Nordborg 








Sanicula elata Buch.-Ham. ex D. 


aHsem 


V 


D.324 


Don 








Sarocococca hooker iana Baill. 


s NP caesp 


I, II 


f.M.,Sch.2083 


Satyrium ciliatum Lindl. 


a G bulb 


V 


By.646,H.64 


5. nepalense D. Don * 


a G bulb 


II, V 


dH.2881,H.69,P.26 


Saussurea auriculata (DC.) Sch. 


aHsem 


V 


By. 622 


Bip.* 








5. candolleana (DC.) Sch. Bip. 


a H scap 


V 


P.R.352 


5. deltoidea (DC.) Sch. Bip. * 


a H scap 


I 


P.s.n. 


5. gnaphalodes (Royle ex DC.) 


a G rad/H ros 


X 


P.G.236, 256, 296 


Sch. Bip. * 








S. gossypiphora D. Don * 


a H scap/G rad 


X 


By.l45,M.1000, 1046, 
1084, P.G.218, 279, 
W.215,Z.1452, Sm.s.n. 


5. graminifolia Wall, ex DC. * 


a H scap/g Ch herb 


X 


M.1061, P.s.n., Z.1450, 




pulv/G rad 




Sm.s.n. 


S. hieracioides Hook.f. 


a G rad/H ros 


X 


P.R.335,W.212,Z.1476, 
Sm.s.n. 


S. leontodontoides (DC.) Sch. 


aHros/Grad/fCh 


VI, IX, X 


By.558, 1135, M.953, 995, 


Bip. 


herb pulv 




1081, W.226, Sm.s.n. 


5. leontodontoides V2cr . filicifolia 


aHros 


IX, X 


P.s.n. (f.Enum) 


(Hook.f.) Hand.-Ma77. 








S. nepalensis Spreng. * 


a H ros 


V 


P.s.n. 


5. simpsoniana (Field. & Gardn.) 


aGrad/fChherb 


X 


M.1046A,S.6583,W.214, 


Lipschitz * 


pulv/H scap 




Z.1516, Sm.s.n. 


5. tridactyla Sch. Bip. ex Hook.f. 


a H scap/G rad 


X 


M.1047,Z.576, Sm.s.n. 


S. uniflora Wall, ex Sch. Bip. 


a H scap/G rad 


III,V 


Beer 25390 


5. wernerioides Sch. Bip. ex 


a f Ch herb pulv 


IX, X 


M.1049,P.G.235,W.217, 


Hook.f. 






Sm.s.n. 


S.yakla C.B.Clarke 


H ros/G rad 


IX, X 


P.R.303,W.385 


Saussurea sp. 


a G rad/ros 


III, IV 


M.919 


Saxifraga andersonii Engl. * 


e g Ch herb pulv 


IX 


D.419, 443, Z.566, Sm.s.n 



PLANTS SOUTH OF MT EVEREST 



257 



S. aristulata Hook.f . & Thorns. 



S. brachypoda D. Don * 



S. brunonis Wall, ex Seringe * 

5. cordigera Hook.f. & Thorns. 
S. diversifolia Wall, ex Seringe 
S. engleriana H. Smith * 
S. filicaulis >yall. ex Seringe 

S. gageana W. W. Smith 
5. georgei Anthony 
5. granulifera H. Smith 

5. hirculoides Decne. 

5. hispidula D. Don 

5. hookeri Engl. & Irmsch. 
S. jacquemontiana Decne. * 

S. lowndesii H. Smith 

5. lychnitis Hook.f. & Thoms. * 
5. cf. melanocentra Franch. * 
5. moorcroftiana (Seringe) Wall. 

ex Sternb. 
5. mucronulata Royle * 

5. mucronulata subsp. 

sikkimensis (Hulten) Hara 
S. nutans Hook.f. & Thorns. 

S. pallida Wall, ex Seringe 



S. palpebrata Hook.f. & Thoms. 
5. parnassifolia D. Don * 

5. perpusilla Hook.f. & Thoms. 

5. pilifera Hook.f. & Thoms. 

5. pseudo-pallida Engl. & Irmsch. 



5. punctulata Engl. 

S.royleiU. Sm.* 

S. saginoides Hook.f. & Thoms. 



S. serrula H. Smith 

5. sibirica L. * 

5. sikkimensis Engl. 



g H herb pulv 


IV, VI 


By.673,H.27,52,M.843, 
907,P.G.214,W.211, 
Z.1479, 1526, 1615B, 
Sm.s.n. 


eHscap 


IV 


Beer 25412, By.489, 666, 
H.7,M.843, Sm.s.n., 
P.s.n.,W.287,Z.1876 


eHros 


VI, IX 


By.487,M.1018,P.R.350, 
W.338, Sm.s.n. 


Hsem 


IV,IX,X 


Beer 10003 


aHsem 


V 


P.48,Z. 1292, 1805 


eHros 


X 


Mc.332, S.7106, Sm.s.n. 


e H caesp/scap/f Ch 


IV, VII 


By.604,M.865,Z.s.n. 


herb pulv 






eHros 


IV, VII, IX 


Beer 9581, M. 1020 


e g Ch herb pulv 


X 


Sw.500 


aHros 


III,V 


dH.2975,M.898,P.R.340, 
W.284,Z.1506, Sm.s.n. 


aHros/fChherb 


IX, X 


By.542,P.G.s.n., Sm.s.n. 


pulv 






a H caesp 


IV, VI, IX 


M.908, 1020,P.G.331, 
W.260 


aHros 


X 


P.47, S.628, W.374 


e H ros/g Ch herb 


IX, X 


M.1086 


pulv 






e H ros/rept/g Ch 


X 


Sw.421 


herb pulv/G rhiz 






a f Ch herb pulv 


VIII-X 


M.1089,Mc.329, Sm.s.n. 


aHros 


X 


By.450, Sm.s.n. 


agChherbpulv/H 


VI 


M. 1026, Z. 1549 


ros 






eHros 


X 


By.443,D.417,M.1013, 
Z.1615a, Sm.s.n. 


e H ros/rept 


X 


BL.2115,Mc.356 


a H ros/Ch herb 


IX 


H.30,W.207, 287, Z. 1526, 


pulv 




1623, Sm.s.n. 


aHsem 


III, IV 


BL.2156, By.515, D.352, 
440, L.2202, Mc.297, 
S.4780 


a H sem 


III, IV 


Beer 9554, D.461 


aHsem 


IV 


By.391,781,M.810, 
Z.1805 


eHros/fChherb 


X 


M. 1064, 1106, Mc.340, 


pulv 




P.G.237, W.232 


e H ros/rept 


VIII 


BL.2143, Sm.s.n. 


aHros 


VIII, IX 


BL.2095,D.398,438, 
M.1012,Mc.333, 
Sm.s.n. 


eHros 


X 


BL.2145,M.1078, 
P.G.274,W.180, Sm.s.n, 


e f Ch herb pulv 


IV, IX 


P.s.n. 


e g Ch herb pulv 


IX, X 


BL.2141,By.778,D.413, 
H.42,M. 1040a, Mc.331, 
P.G.238,W.161,223, 
277, Sm.s.n. 


e H rept/caesp 


IV, V 


P.50 


a H ros/sem 


IV 


B.5777,By.512 


aHscap 


III, IV 


Beer 25512, S. 1644 



258 

S. stella-aurea Hook.f. & Thorns. 

5. stenophylla subsp. hoffmeisteri 

(Klotzsch) Hara * 
5. strigosa Wall, ex Seringe * 

Schisandra grandiflora (Wall.) 

Hook.f. & Thorns. * 
Scopolia stramonifolia (Wall.) 

Shrestha * 
Scrophularia pauciflora Benth. 
Sedum chauveaudii R.-Hamet 
5. gagei R.-Hamet 
S. oreades (Decne.) R.-Hamet * 
5. triactina Berger 
5. trullipetalum Hook.f. & 

Thoms. 
Selinum candollii DC. 
S. cortioides C. Norman 
5. tenuifolium Wall, ex C. B. 

Clarke * 
Senecio alatus Wall, ex DC. 
S. albopurpureus Kitam. 
S. cappa Buch.-Ham. ex D. Don 
S. chrysanthemoides DC. * 
S. diver sifolius Wall, ex DC. 
S. graciliflorus DC. * 
5. lancifera J. R. Drummond 
S. rufinervis DC. 
5. scandens Buch.-Ham. ex D. 

Don* 
5. triligulatus Buch.-Ham. ex D. 

Don 
S. wallichiiDC. 
Sibbaldia cuneata Homem. ex 

Kuntze * 
5. micropetala (D. Don) 

Hand.-Mazz. 
S. perpusilloides (W. W. Smith) 

Hand.-Mazz. 
5. purpurea Royle * 

Silene aff. gonosperma (Rupr.) 

Bocquet 
S. gonosperma subsp. 

himalayensis (Rohrb.) Bocquet 

var. himalayensis 
S. nigrescens (Edgew.) Majumdar 
S. setisperma Majumdar 
5. thomsonii Majumdar 
Sinocarum sp. 
Skimmia laureola (DC.) Sieb. & 

Zucc. ex Walp. * 
Smilacina oleracea (Baker) 

Hook.f. * 
5. purpurea Wall. * 
Smilax menispermoidea DC. 
Sorbus cuspidata (Spach) 

Hedlund * 



G. MIEHE 



e H ros/rept/f Ch 


X 


BL.2145,M.1100,Sm.s.n. 


herb pulv 






e H ros/rept 


X 


M.1118 


a H sem/rept 


IV 


By.660,M.906,P.51, 
Z.1660 


a d PL frut 


I 


Z.716 


a H scap 


V 


f.M. 


a G rhiz/scap 


III 


B.5679 


a Ch 1 succ 


IV 


Z.1679 


a H scap 


IV 


H.22,M. 1020, Z. 1609 


a Ch 1 succ 


VI, IX 


By. 620 


a Ch 1 succ/caesp 


I, IV 


Mc.270 


a Ch 1 succ 


IV 


M.862, 960, W.273, 301 


aGrad 


? 


By.13,245,655 


aG rad 


VI, IX 


M.lOOla 


aGrad 


V 


M.848, 883a 


aHsem 


I, II 


Z.1634, 1770 


aHros 


VIII, X 


M.1052 


a Ch suff 


II 


Z.1740A, 1845 


a H sem/scap 


II, V 


By. 197, 399 


a H scap 


II, V 


M.827, P.s.n. 


a G rad/scap 


II, V 


Z.1760 


aHsem 


v' 


M.829 


aHsem 


? 


P.s.n. 


a Ch suff/L 


ii 


dH.2909, Sch.2363, W.383, 
Z.1740 


a Ch suff 


n 


dH.2901,S.4487 


aHsem 


III,V 


Z.1662 


a g Ch herb pulv/G 


V 


Z.658, 695 


rhiz 






aHsem 


V 


BL.2171,D.346,Z.789 


aGrhiz/fChherb 


VII, X 


Mc.379 


pulv 






aGrhiz/fChherb 


IX, X 


M. 1039, 1077, Mc.377, 


pulv 




Z.568, Sm.s.n. 


a H caesp 


VII 


M.1027 


a H caesp 


IX 


BL.2187, By.432, Sm.s.n. 


a G rhiz/H sem 


VI, VII 


M. 1006, Sm.s.n. 1 


a H scap 


? 


Sm.s.n. 


a H sem 


ill, IV 


M.968, P.s.n. 


aGrad 


VI 


M.s.n. 


s NP caesp 


I 


S.2064, 4477 


a G rhiz 


n 


Mc.267, S.4627 


a G rhiz 


II, III 


Mc.291 


s d PL frut 


I 


Mc.55, 423, 1783 


a Mes P scap 


III 


M.824 



PLANTS SOUTH OF MT EVEREST 



259 



5. foliolosa (Wall.) Spach 


a Mi P scap 


m 


M.893 


5. hedlundii Schnexd. 


a Mi P scap 


I 


P.s.n. 


S. microphylla Wenzig * 


a Mi P scap 


m 


By.7, 839, 1155, D.451, 
S.6580, 7143, 7147 


5. ursina (Wenzig) Decne. 


a Mi P scap 


m 


Mc.276 


Soroseris hookeriana (C. B. 


aHros 


X 


M.998, W.254 


Clarke) Stebb. * 








S. hookeriana (C. B. Clarke) 


aHros 


X 


P.G.247, Sm.s.n. 


Stebb. subsp. hookeriana 








S. pumila Stebb. 


a H ros 


X 


By.382,P.G.217,Sm.s.n. 


Spathoglottis ixioides (D. Don) 


obi. a E Ch succ 


IV 


By.366, M.806, L.016 


Lindl. * 








Spiraea arcuata Hook.f. * 


a NP caesp 


III,V 


By.439,M.933, P.s.n., 
Sm.s.n. 


5. bella Sims 


a NP caesp 


II 


S.4628 


5. canescens D. Don * 


a NP caesp 


n 


S.7122 


Spiranthes sinensis (Pers.) Ames 

* 


aGrad 


V 


H.62-64, M.835 


Stellaria congestiflora Hara 


a H rept 


VIII, X 


W.246 


5. decumbens Edgew. * 


e H rept/e f Ch herb 
pulv 


X 


M.1022, Sm.s.n. 


5. decumbens var. polyantha 


a H rept 


X 


BL.2028, By.387, M.971 


Edgew. & Hook.f. 








5. decumbens var. pulvinata 


a f Ch herb pulv/a 


VIII, X 


M.s.n. 


Edgew. & Hook.f. 


Hrept 






5. graminea L. 


a H rept 


IV, VIII 


M.897 


S. lanata Hook.f. ex Edgew. & 


a H rept 


? 


W.373 


Hook.f. 








S. sikkimensis Hook.f. ex Edgew. 


Trept 


? 


W.380 


& Hook.f. 








Streptopus simplex D. Don * 


aGrad 


m 


By.l38,BL.2017,L.0158, 
Z.777 


Sunipia bicolor Lindl. 


obi. e E Ch succ 


I 


Sch.2312 


Swertia cuneata D. Don 


aHsem 


VII, VIII 


M.949, 1057,P.160,R.328 


5. rfitoam C.B.Clarke 


a H scap 


V 


M.918 


5. /iooA:en C.B.Clarke* 


a H sem/G rad 


VII 


S.7087 


5. kingii Hook.f. 


a H sem/G rad 


V 


L.0419, W.290 


5. multicaulis D. Don 


a G rad/H caesp 


VII 


By.504, 524, D.339, H.44, 
P.G.209,G.285,W.234, 
Sm.s.n. 


Sympagis petiolaris (Nees) 




II 


Z.1743 


Bremek. 








Symplocos paniculata (Thunb.) 


s/a Mi P scap/caesp 


I 


S.4623 


Miq.* 








5. sumuntia Buch.-Ham. ex D. 


s Mi P scap 


I 


S.4483 


Don 








Tanacetum atkinsonii (C. B. 


aHsem 


V,VI 


P.s.n. 


Clarke) Kitam. * 








T. gossypinum Hook.f. & Thoms. 


a H ros/scap 


IX, X 


By.ll76,H.39, Sm.s.n., 


ex C.B.Clarke* 






M.1031,1070,P.G.211, 

G.295,Sch.2315,W.179, 

Z.1509 


Taraxacum mitalii van Soest 


a G rad/H ros 


V-VII 


Z.270 


T. monochlamydeum 


a G rad/H ros 


V 


Z.1763 


Hand.-Mazz. 








T. pseudostenoceras van Soest 


a G rad/H ros 


V,VI 


Z.1603 


Taxus baccata subsp. wallichiana 
(Zucc.) Pileer 


b N/Mi P scap 


I 


f.M.,Sch.2056,2939, 
S.4496 



260 

Thalictrum alpinum L. * 

T. cultratum Wall. * 

T. elegans Wall, ex Royle 

T. reniformeWall. * 

T. virgatum Hook.f. & Thorns. * 

Thermopsis barbata Royle * 



Thesium himalense Royle ex 

Edgew. 
Tiarella polyphylla D. Don 
Tofieldia himalaica Baker 
Trachydium aff. royleilJmdX. + 
Trigonotis multicaulis (DC.) 

Benth. ex C.B.Clarke 
T. ovalifolia (Wall.) Benth, ex C. 

B. Clarke 
T. rotundifolia (Wall, ex Benth.) 

Benth. ex C.B.Clarke* 
Triosteum himalayanum Wall. 
Triplostegia glandulifera Wall, ex 

DC. 
Tripogon filiformis Nees ex 

Steudel 
Tripterospermum volubile (D. 

Don) Hara 
Trisetum scitulum Bor 
T. spicatum (L.) K. Richter 
Trollius pumilus D. Don 
Tsuga dumosa (D. Don) Eichler * 
Typhonium diversifolium Wall. 

ex Schott * 

Urtica dioica L. 
Utricularia striatula Sm. 

Vaccinium dunalianum Wight 
V. glauco-album Hook.f. ex C. B. 

Clarke 
V. nummularia Hook.f. & 

Thorns, ex C. B. Clarke * 
V. retusum (Griff.) Hook.f. ex C. 

B. Clarke 
Valeriana hardwickii Wall. 

V. jatamansii Jones * 
V. stracheyi C. B. Clarke 
Verbascum thapsus L. 
Veronica cana Wall, ex Benth. 
V. cephaloides Pennell 

V. lanuginosa Benth. ex Hook.f. 

* 

V. himalensis D. Don 
V. umbelliformis Pennell 
Viburnum erubescens Wall, ex 

DC* 
V. grandiflorum Wall, ex DC. * 



G. MIEHE 

a H ros VI, VII, IX 



aHsem 
aHsem 
aHsem 
aHsem 

aGrad 



aGrad 



V 
V 
V 

I, II 



V,VII 



D.394,400,BL.2111, 

Z.644, Sm.s.n. 
S.7140,Sm.s.n. 
P.s.n.,W.405 

By.l76, 191,M.890,S.7146 
BL.2169,D.323,P.s.n., 

Sch.2099 
By.511,552,949,dH.2956, 

Sch.2053, 2054, 2174, 

Z.1646 
P.42,Z.501,682 



aHsem 


I, II 


P.53,Z.448,768 


a G rhiz 


VI, VII 


M.964 


aGrad 


? 


By.561 


a H caesp 


ill, IV 


Beer 10016, S.653 


a H scap 


ni 


Mc.357, S.7088 


a H caesp 


V,VII 


BL.2204, S.652 


a H scap 


I, III, V 


L.0654,P.s.n.,Z.679 


aGrad 


7 


H.12,W.364 


a H caesp 


IV 


M.808, 817c 


aelHL 


I, II 


Sch.2068 


a H caesp 


VI, VII 


P.G.303 


a H caesp 


VI 


By. 1071, M. 1021a, P.R.349 


aHros 


VIII 


BL.2080 


b Mes P scap 


I 


f.M.,Sch.2340,2360 




I 


By.225 


a G rhiz 


V 


f.M. 


a hyd H rept 


IV 


P.267 


s E Ch frut 


I 


BL.2021 


s NP caesp 


I 


Sch.2163 


s fac E Ch frut rept 


I 


S.4605,Z.266,382 


fac E/s NP caesp/Ch 


I 


P.B. 75,8.4475 


frut 






a H scap 


II 


By.320, D.358, Sch.2077, 
Z.1672 


aHsem 


n 


Z.263 


a H scap 


n 


Z.1317 


aHsem 


V 


By.l82,Sch.2061,P.56,57 


a H scap 


I 


P.260 


a H scap 


? 


W.191 


a H rept/G rhiz 


X 


BL.2176,M.1079,Mc.375, 
Sw.88-99 


a H scap 


? 


By. 130 


a H scap 


? 


W.308 


a Mi P caesp 


I, II 


Z.734 



a NP caesp 



m 



Z.405 



PLANTS SOUTH OF MT EVEREST 



261 



V. nervosum D. Don * 
Vincetoxicum hirundinaria subsp. 

glaucum (Wall, ex Wight) Hara 

* 


a Mi P caesp 
a G rad/scap 


III 
V 


Mc.35, S.4523, Z.795 
Sch.2249, Z.675, 714 


Viola biflora L. * 

V. pilosa Blume 

V. thomsonii Oudemans 


aHros 

aHros 
aHros 


II, III, V 

I, III 
I 


By.ll0,W.366,Z.579,652, 

702, 1774, Sm.s.n. 
Onyon 10, S.4479, Z.397 
S.4486 


Waldheimia glabra (Decne.) 
Regel * 


a H ros/G rhiz 


VIII, X 


By.448, H.36, dH.2999, 
M.973,Mc.315,P.R.345, 
W.174,181,Z.1453, 
Sm.s.n. 


Youngia japonica (L.) DC. 

y. racemifera (Hook.f.) Babcock 

& Stebb. 
Ypsilandra yunnanensis W. W. 

Smith & Jeffrey 


aHsem 

a H sem/a G rad 

aGrad 


I, II 
III 

V,VII 


Z.748 
M.976,W.319 

Beer 25356 


Zanthoxylum nepalense Babu * 
Z. oxyphyllum Edgew. * 


a/s NP caesp 
a d PL frut/NP 
caesp 


II 
II 


Z.758 

BL.2015,Z.465, 1695, 1866 



Alphabetical list of ferns and fern allies 

Adiantum pedatum L. 
Anogramma leptophylla (L.) 

Link 
Araiostegia pulchra (Don) Copel. 
Arthromeris himalayensis 

(Hook.) Ching 
A. wallichiana (Sprengel) Ching 
Asplenium ensiforme Wall, ex 

Hook. & Grev. 
A. varians Wall. 
Athyrium drepanopterum 

(Kunze) A. Br. 
A. duthiei (Bedd.) Bedd. 
A. fimbriatum (Wall.) T. Moore 
A. mackinnoni (Hope) 

Christensen 
A. pectinatum (Wall, ex Mett.) 

Bedd. 
A. schimperi Moug. ex Fee 
A. schizochlamys Ching 
A. spinulosum (Maxim.) Milde 

Cheilanthes albomarginata C. B. 

Clarke 
C. farinosa (Forsk.) Kaulf. 
C. subvillosa Hook. 
Coniogramme affinis Hieron. 
C. caudata (Wall, ex Ettingsh.) 

Ching 
C. intermedia Hieron. 
Crypsinus erythrocarpus (Mett. 

ex Kuhn) Tagawa 



erhiz 


III,V 


Z.1688, 1779 


a caesp 


I 


Z.1754 


a rhiz 


II 


Z.674 


a rhiz 


I 


dH.2888, Z.256 


a rhiz 


I 


Z.1871 


e caesp 


I, II 


Z.1896 


a caesp 
a/e caesp 


III 

I 


W.357 
Z.1019 


a rhiz 


III 


Z.646 


a/e caesp 
a caesp 


I 
I, III 


Z.1870 
Z.1682 


a rhiz 


I, III 


Z.1665 


a rhiz 


I 


Z.678, 1752 


a caesp 


V 


M.885A, 912A, 927 


a rhiz 


m 


Z.677, 1685, 1794 


a caesp 


I 


Z.761 


a caesp 
a caesp 
erhiz 


? 

V 
IV 


W.354 
Z.1823 
Z.725 


erhiz 


IV 


Z.454 


erhiz 


IV 


Z.1749 


erhiz 


I, III 


Z.1689B 



262 

Cryptogramma brunoniana Wall. 

ex Hook. & Grev. 
Cystopteris dickieana Sim 

C. filix-fragilis (L.) Bernh. 

Drynaria mollis Bedd. 
Dryopteris acuto-dentata Ching 

D. barbigera (T. Moore ex 
Hook.) Kuntze subsp. 
barbigera 

D. chrysocoma (Christ) C. Chr. 
D.fibrillosa (C. B. Clarke) 

Hand.-Mazz. 
D. marginata (Wall, ex C. B. 

Clarke) Christ 
D. serratodentata (Bedd.) Hayata 
D. sublacera Christ 
D. wallichiana (Spreng.) N. 

Hylander 

Equisetum diffusum D. Don 

Gymnocarpium oyamense (Bak.) 
Ching 

Lepisorus clathratus (C. B. 

Clarke) Ching 
L. contortus (Christ) Ching 
L. kashyapii (Mehra) Mehra 
L. loriformis (Wall.) Ching 
L. variabilis Ching & S. K. Wu 
Leucostegia delavayi (Bedd.) 

Ching ex Christensen 
Lycopodium clavatum L. 
L. veitchii Christ 

Mecodium polyanthos (Sw.) 
Copel. 

Notholaena marantae (L.) Desv. 



Oleandra wallichii (Hook.) C. 

Presl 
Onychuum contiguum Hope 
Osmunda claytonia var. vestita 

(Wall.) Milde. 

Phymatodes crenatopinnata (C. 

B. Clarke) Ching 
P. ebenipes (Hook.) Ching 
P. malacodon (Hook.) Ching 
P. stewartii (Bedd.) Ching 
Plagiogyria communis Ching 
Polypodium amoenum Wall, ex 

Mett. 
P. microrhizoma C. B. Clarke ex 

Baker 



G. MIEHE 








a caesp 


III,V, 


VII 


M.986,W.256,Z.1562 


e caesp 


VII 




M.1010A,Z.1618 


a caesp 


V 




Z.639, Sm.s.n. 


arhiz 


I, IV, 


V 


Z.1639 


a caesp 
a caesp 


V 
VI 




M.885B,961 
M.886 


a caesp 
a caesp 


VI 
III, VI 


Z.1642, 1684 
Z. 1681, 1776 


a caesp 


I, III, 


IV 


Z.273 


a caesp 


VII 




M.1010B,Z. 1542, 1567 


a caesp 
e caesp 


I 
I, III 




Z.385, 488 
Z.455, 1832 


a G rhiz 


I, III 




W.430 


arhiz 


I 




Z.719 


e/a rhiz 


m 




Z.507, 638 


e rhiz 


III 




Z.377, 396 


erhiz 


I 




Z.721 


erhiz 


III 




Z.791B 


e/a rhiz 


m 




W.345 


arhiz 


V 




Z.1625 


b Ch herb rept 
b Ch herb rept 


I 
? 




W.387, Z.427, 1890 
W.352 


arhiz 


I, III, 


V 


Z.703, 1895 


a caesp 


I, III- 


-V 


dH.2965,M.840,885c, 
925, Z.265, 629, 1650, 
1713 


erhiz 


IV 




Z.754 


arhiz 


11, V 




dH.2882,Z.1658 


e caesp 


II 




Z.715 


erhiz 
erhiz 


I, III 
III 




Z.\6%9B {CandollealS: 

206 [1956]) 
Z.715B, 1689 


erhiz 


III, VII 


W.298, Z.637, 1334, 1624 


erhiz 


I 




Z.738 


e caesp 
e/a rhiz 


IV 

I, III, 


,IV 


Z.257, 460 
Z.753 



erhiz 



M.912B 



Polystichum acanthophyllum 

(Franchet) Christ 
P. discretum (Don) Diels 
P. lachenense (Hook.) Bedd. 
P. mehrae Fraser- Jenkins & 

Khullar 
P. neolobatum Nakai 
P. nepalense (Spreng.) C. Chr. 
P. shensoense Christ 
P. stimulans C. Presl 
P. thomsonii (Hook.f.) Bedd. 

Pteridium aquilinum (L.) Kuhn 

Pteris cretica L. 

P. dactylina Hook. 

P. subquinata Wall, ex Hope 

P. walUchianaJ. Agardh 

Selaginella chrysocaulos (Hook.f. 
& Grev.) Spring 



PLANTS SOUTH OF MT EVEREST 
e caesp I, III 



263 



Z.384 



a caesp 


I 




Z.455 


e caesp 


VI-X 




M.lOlOcZ. 1449, 1595 


e caesp 


II 




N.9 


e caesp 


I 




Z.1680 


a caesp 


I, III, 


IV 


Z.1742 


a caesp 


III 




M.975 


e caesp 


IV 




Z. 1666, 1737 


a caesp 


IV 




W.356, Z.399, 747, 826b, 
1736 


arhiz 


II, III 


,V 


M.s.n. 


e rhiz 


I 




Z. 1738, 1847 


erhiz 


I 




Z.267, 487, 1737 


e caesp 


IV 




Z.1826 


e caesp 


IV 




Z.1873 



Z.1735 



Thely pteris levingei (C. B. 

Clarke) Ching 
Tricholepidium normale (D. 

Don) Ching 


arhiz 
erhiz 


m 

IV 


Z.1686 
Z.1750 


Vittariaflexuosa Fee 


erhiz 


IV 


Z.1748 


Woodsia andersonii (Bedd.) 
Christ 


a caesp 


? 


W.355 


W. elongata Hook. 
W. lanosa Hook. 


a caesp 
a caesp 


IV 

? 


Z.680, 1751,1775,1809 
W.358 



Acknowledgements 

In the summer and autumn 1982 I had the opportunity to accompany Professor Dr Matthias Kuhle, 
Department of Geography, University of Gottingen, during a three months expedition to the Khumbu 
Himal. I have to thank him first for giving me the chance to carry out field work and to collect plants. The 
expedition was sponsored by the DFG. My own small collection of plants was determined during a four 
weeks stay at BM in December 1982 and January 1983 and in March and April 1985, supported by the 
British Council, by the kind permission of the Keeper of Botany, Mr J. F. M. Cannon, and the Head of the 
General Herbarium, Mr A. O. Chater, to whom I am highly indebted. Only by the help of all members of 
staff and the clearly arranged unique collection of the herbarium I was able to check all life forms of plants 
collected in the Khumbu Himal and the adjacent valleys. In many questions I was able to profit of the 
experience of Mr J. D. A. Stainton. Special thanks are also extended to Dr E. Launert for his help in every 
situation, and to Mr J. F. M. Cannon and Mr J. R. Laundon for their considerable editorial efforts. 



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268 G. MIEHE 

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Bichsel, H. Kienholz). 



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Further genera of the Biddulphiaceae 
(diatoms) with interlocking hnking 
spines 

Robert Ross & Patricia A. Sims 



Botany series Vol 16 No 4 26 November 1987 



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Further genera of the Biddulphiaceae (diatoms) with 
interlocking linking spines 

Robert Ross and Patricia A. Sims 

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

Contents 

Synopsis 269 

Introduction 269 

Sources of specimens and records 270 

Terminology 271 

Taxonomic account 

I. Solium Heiberg 271 

II. Pseudorutilaria (Grove & Sturt ex De Toni & Levi) 

Grove & Sturt ex De Toni 276 

III. Mom/e R. Ross & P. A. Sims 284 

IV. Mfl/wma R. Ross & P. A. Sims 285 

V. fionea R. Ross & P. A.Sims 286 

VI. DextradonatorK. Ross & P. A. Sims 288 

Discussion 289 

Acknowledgements 292 

References 292 

Description of plates 295 

Plates 299 

Synopsis 

Descriptions are given of a number of genera of the Biddulphiales, all except one with interlocking linking 
spines. Solium Heiberg, a monotypic genus, is distinguished from Trinacria. Pseudorutilaria (Grove & 
Sturt ex De Toni & Levi) Grove & Sturt ex De Toni is close to Solium but has some unique characteristics. 
Five new species from the Eocene of the southern oceans are described. Monile R. Ross & P. A. Sims, a 
monotypic genus from the Miocene of southern France with similarities to both Pseudorutilaria and 
Biddulphia, is described as new. A new genus, Maluina R. Ross & P. A. Sims, is proposed for Hemiaulus 
centralitenuis R. Ross & P. A. Sims, and another monotypic new genus, Bonea R. Ross & P. A. Sims, is 
established for a tripolar diatom from the Eocene of the U.S.S.R. close to Strelnikovia R. Ross & P. A. 
Sims. New information about Dextradonator eximius (Grunow) R. Ross & P. A. Sims is presented. The 
position of genera within the family Biddulphiaceae is discussed but presents some difficulties, especially 
because of possible parallel evolution. The limits of the order Biddulphiales are also considered. 

Introduction 

In a recent paper (Ross & Sims, 1985) we dealt with a group of related genera in the diatom 
family Biddulphiaceae characterized by the possession of interlocking linking spines and well 
developed pseudocelli. In the course of preparing this we examined all the other members of the 
family with interlocking linking spines but concluded that none of these were close to the group 
with which we were dealing there. They belonged to four genera and our studies of these with the 
scanning electron microscope produced new information recorded here. One of these genera is 
Solium Heiberg (1863), for our examination of 5. exsculptum Heiberg, its type species, showed 
that it should not be placed in Trinacria Heiberg (1863), the genus to which it was transferred by 
Hustedt (1930). Another is Pseudorutilaria (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex 
De Toni (1894), of which we found five undescribed species in Eocene material from the 



Bull. Br. Mus. Nat. Hist. (Bot.) 16 (4): 269-311 Issued 26 November 1987 



270 R. ROSS AND P. A. SIMS 

Falkland Plateau in the South Atlantic. This genus is apparently close to Solium, but it has 
remarkable structural features on its central portion that clearly distinguish it. We re-examined 
the diatom that we earlier described as Hemiaulus centralitenuis R. Ross & P. A. Sims (in Ross, 
Sims & Hasle, 1977) and concluded that it could not be retained in Hemiaulus Heiberg (nom. 
cons, prop., see Ross, 1985), but that it was necessary to establish for it a new genus, which we 
have called Maluina. The fourth genus with interlocking linking spines is DextradonatorR. Ross 
& P. A. Sims (1980), and we present here additional information about D. eximius R. Ross & P. 
A. Sims. 

We also describe here another new genus, Bonea, based on a single new species with 
interlocking linking spines. This, which comes from the Eocene of the U.S.S.R., has many 
similarities with the genera we described in our previous paper, especially Strelnikovia R. Ross 
& P. A. Sims (1985), but it had not come to our notice at the time when that paper was prepared. 
Another new genus described here, Monile, is also based on a single new species, one that comes 
from the Miocene of southern France. It does not have interlocking linking spines, but it is 
included here because, although there is little to separate it from Biddulphia Gray (1821), it has 
many features in common with Pseudorutilaria, as well as some in which it resembles Kera- 
tophora Pantocsek (1889) and Strelnikovia R. Ross & P. A. Sims. Its affinities are puzzling. 



Sources of specimens and records 

In the distribution lists for the species considered in this paper there are many records taken 
from literature, and all the information available about these is to be found in the references 
given. We have endeavoured to give a complete account of the known distribution of the species 
dealt with, but we realize that our coverage of Russian literature is incomplete, although 
probably sufficient to give a reasonably adequate picture. Many of the specimens that we have 
ourselves examined come from localities of which we gave details in our recent paper (Ross & 
Sims, 1985). The additional localities from which we have seen specimens are discussed below. 

U.S.S.R. 

Northern Urals 

One of the samples that the British Museum (Natural History) received from Mr Brigger was 

labelled 'N. Urals, U.S.S.R., Oligocene'. We have no further information about this material 

but presume that it is from one of the samples that are the basis of the records from the lower 

Oligocene of the northern and central Urals in Diatomovyi Analiz (Proshkina-Lavrenko et al., 

1949a). 

Germany 

Greifswalder Oie, Rostock 

This is one of the localities from which Schulz (1927) reported lower Eocene diatoms. The 

specimen from this locality that we have seen was mounted by Hustedt and came from Schulz's 

sample. 

Denmark 

All the specimens from Denmark that we have examined come from the Moler formation of 
lower Eocene (Ypresian) age. The precision with which the locality is indicated varies; the 
specimens are said to come from: Jutland, Glyng0re, Skive, Limfjord, Island of Mors, 
Nykjobing, Ejerslev, Island of Fur. 

France 

Saint Laurent-la Vernede, Gard 

The Miocene deposit from this locality is described by Lefebure (1935). We are not aware of any 

more precise information about its geological age. 



GENERA OF THE BIDDULPHIACEAE 271 

Barbados 

We have examined specimens from two localities in Barbados additional to those from which we 
reported specimens in our previous paper (Ross & Sims, 1985). These are Bissex Hill and 
Malvern Hill. No information about the level of these samples within the Oceanic Beds, which 
range from middle Eocene to lower Miocene, is available to us. 

'Lament E 8' 

This is the only label on one sample received from Mr Brigger. Neither the California A5:ademy 
of Sciences nor the Lamont-Doherty Geological Observatory have any record giving locality 
data for this. Its content suggests very strongly that it comes from the middle to upper Eocene of 
the Falkland Plateau area. Mr Brigger received a number of such samples from the Lamont- 
Doherty Geological Observatory and ii is presumably one of these. 

Terminology 

The terminology used in the descriptions of the genera and species is that proposed by Ross et al. 
(1979), supplemented by terms that we adopted in our most recent paper (Ross & Sims, 1985: 
285) and one proposed by von Stosch & Simonsen (1984: 11), viz. 'interstrial pores'. In addition 
we use here two new terms, 'subocellus' and 'pseudo-antiligula'. 

In Solium Heiberg, Pseudorutilaria (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De 
Toni, and Monile R. Ross & P. A. Sims there is a cluster of perforations through the valve on the 
distal side of the upper parts of the elevations; in Monile this also extends onto the summit of the 
elevations (Fig. 64). Vela are present in these in Solium (Fig. 5) and Pseudorutilaria (Fig. 42), 
and probably also in Monile. There is, however, no transition between the small areolae in these 
clusters and the areolae on the rest of the valve, and in this they differ from the pseudocelli of 
such genera as Biddulphia S. F. Gray (Ross & Sims, 1971: pi. 1 fig. 2), Briggera R. Ross & P. A. 
Sims (1985: pi. 5 fig. 6), and Trigonium Cleve (Ross & Sims, 1971: pi. 2 fig. 6). On the other hand 
there is no thickened ring of silica around these clusters of small areolae such as surrounds the 
ocelU oi Amphitetras Ehrenberg (Ross & Sims, 1971: pi. 5 fig. 2) and the other genera included in 
the Eupodiscaceae and the ocelluli of the Cymatosiraceae (Hasle et al. , 1983). For these clusters 
of small areolae with a definite but unthickened margin we use the term 'subocellus'. We have 
also used this term for the small cluster of more tightly and uniformly packed perforations on the 
distal side of the tip of the elevations in Maluina R. Ross & P. A. Sims (Fig. 68) as this cluster 
also has no thickened rim; it may well be, however, that this cluster consists of porelli, not 
areolae. 

In Maluina centralitenuis (R. Ross & P. A. Sims) R. Ross & P. A. Sims there is at each pole of 
the valve a downward projection of the mantle, i.e. one parallel to the pervalvar axis. Similar 
projections are present on the girdle bands of many species and are termed 'antiUgulae'. This 
term cannot, however, be used for structures that are part of the valve; hence we use for them 
the term 'pseudo-antiligula'. 

Taxonomic account 

I. SOL/t/M Heiberg, Consp. crit. Diat. Dan.: 52 (1863). 

Hemiaulus subgen. Solium (Heiberg) Van Heurck, Treat. Diat., transl. Baxter: 455 (1896). 

Frustules united in short, straight, inseparable chains, the pervalvar axis from about half to 
about twice as long as the diameter of the valve. Valves multipolar, with short projections at 
each pole cut off from the central portion by transverse sulci or internal costae. Elevations at 
each pole moderately stout, not expanded above, with a subocellus on the distal side near the 
tip, linking spines on the proximal side of the tip expanded above and interlocking, except those 
of the end cells of chains. A well developed marginal ridge between the elevations. Mantle with 
the upper part areolate and with anastomosing external costae that continue onto the elevations, 
the lower part hyaline and deeply concave. Areolae irregularly scattered on the central portion 
of the valve, decussate or irregularly scattered on the projections, each occluded by a vola. 



272 R. ROSS AND P. A. SIMS 

Scattered hollow spines or occluded processes sometimes present on the central portion of the 
valve. One labiate process on the central portion of the valve near the base of a projection, its 
internal opening sessile and straight, its external part tubular. 

Type species: Solium exsculptum Heiberg (loc. cit.) 

Heiberg (1863) established the genera Trinacria and Solium for tripolar and quadripolar 
diatoms with the frustules united in chains by elevations with Unking spines. Bipolar diatoms 
forming similar colonies he placed in Hemiaulus. Hemiaulus Ehrenberg (1844) has been shown 
to be a later synonym of Eucampia Ehrenberg (1839), and the conservation of Hemiaulus in the 
sense in which Heiberg and all subsequent authors have used it has recently been proposed 
(Ross, 1985). H. L. Smith (1872) did not consider that the number of poles was a sufficient basis 
for distinguishing genera, and he regarded Trinacria and Solium as synonyms of Hemiaulus, as 
did Van Heurck (1896), who treated Trinacria and Solium as subgenera of Hemiaulus, and 
Schiitt (1896), who was the only one of these authors to publish any new combinations under 
Hemiaulus. Most authors, however, maintained the three genera as separate until Hustedt 
(1930) united Trinacria and Solium under the name Trinacria, a genus he regarded as separate 
from Hemiaulus. 

We would agree with the view that the number of poles is not by itself a sufficient basis for 
generic separation, and we are not here concerned with the question of whether or not there are 
other differences on which to base a separation of Trinacria from Hemiaulus. The differences, 
however, between Solium exsculptum Heiberg and Trinacria regina Heiberg, the type species of 
the two genera, are such as to justify placing them in separate genera. In Trinacria regina the 
linking spines are never expanded above and interlocking, there is no subocellus but only a 
poorly developed pseudocellus, the mantle is vertical throughout, and the areolae are occluded 
by cribra; in Solium exsculptum Xhe, linking spines are normally expanded above and interlock- 
ing, there is a well developed subocellus, the lower part of the mantle is strongly concave, and 
the areolae are occluded by volae. We attach particular importance to the presence and nature 
of the subocellus and the concavity of the lower part of the mantle as features separating Solium 
from Trinacria because they suggest, as we argue below (p. 291), that Solium may be more 
closely related to such members of the Eupodiscaceae as Eupodiscus J. W. Bailey (1851), 
Triceratium Ehrenb. (1839) sensu stricto, and Amphitetras Ehrenb. (1840), than to Biddulphia 
Gray (1821) or Hemiaulus Heiberg (1863). Trinacria, on the other hand, is much closer to 
Hemiaulus Heiberg and the majority of genera of the Biddulphiaceae. 

The genus to which Solium is most closely related is Pseudorutilaria (Grove & Sturt ex De 
Toni & Levi) Grove & Sturt ex De Toni and we discuss their resemblances and differences after 
our description of that genus. 

Solium exsculptum Heiberg is the only known species of the genus. The two others that have 
been referred to it, S. jeremiae (Truan & Witt) De Toni (1894) and S. princeps M. Peragallo (in 
Tempere & Peragallo, 1912) are both quadripolar species of Trinacria. S. jeremiae was originally 
described as Trinacria jeremiae Truan & Witt (1888), but S. princeps cannot be called Trinacria 
princeps as that name is pre-occupied. It has, however, been described independently as a new 
species, Trinacria cristata Goxahos {\9^2) . 

I 
1. Solium exsculptum Heiberg, Consp. crit. Diat. Dan. : 52 (1863). 
(PI. 1, PI. 12 figs 86-88) 

Hemiaulus exsculptus (Heiberg) Schiitt in Engler & Prantl, Naturl. Pflanzenfam. 1 (lb): 97 (1896). 
Trinacria exsculpta (Heiberg) Hustedt in Rabenhorst, Krypt.-Fl Deutschl, Osterr. u. d. Schweiz 1 (1): 
889 (1930). 

Valves with 4 or 5 projections separated from the central portion by deep sulci; length of side in 
4-polar specimens 30-110 /oim, maximum diameter of 5-polar specimens 35-45 /am, height at 
centre of valve 7-5-16 jxm; central portion with straight sides between the projections, weakly ^ 
domed; projections semi-circular to narrowly triangular with obtuse apices, slightly domed and 
rising towards the elevations. Elevations arising at the apices, not expanded above; height to 



GENERA OF THE BIDDULPHIACEAE 273 

their tops 18-38 ^im. Areolae poroid, 0-3-0-7 fxm in diameter, occluded by volae, with a raised 
external rim from which one or two very short cylindrical spines sometimes arise, irregularly 
scattered and 5-6 in 10 fxm on the central portion of the valve, the projections, the elevations, 
and the upper part of the mantle, or decussate and rather closer, 7-8 in 10 fxm, on the 
projections. Sparse scattered hollow spines or occluded processes sometimes present on the 
central portion of the valve of end cells. Subocelli on the upper part of the distal face of the 
elevations, circular and c. 4 /am in diameter, surrounded by a narrow hyaline unthickened rim, 
their areolae in radial rows, c. 32 in 10 /am. 3-6 linking spines on the proximal sides of the tips of 
the elevations, usually expanded above and interlocking with those of the sibling valve, 
sometimes (on end cells) tapering upwards, c. 4 /am long; c. 6 short spines c. 0-5 /am tall on the 
distal side of the tips of the elevations. One labiate process on the central portion of the valve 3-5 
/am proximal to a sulcus, opposite the centre of the sulcus or, more frequently slightly offset in a 
clock- wise direction when the valve is viewed from its external side, its inner opening parallel to 
the sulcus, its outer tube almost vertical to strongly incUned distally with its tip touching the 
surface of the adjacent projection. 

forma exsculptum (Figs 1-3, 5, 6, 86, 88) 

Valves quadripolar. 

Paleocene. Indian Ocean, 14° 46-7' S. , 88° 54-4' E. , 4780 m depth. Vityaz station 6744-40 (Mukhina, 1974, 
1976). 

Cape basin. South Atlantic Ocean, 29° 29-055' S., 3° 30-74' E., 4805 m depth. Deep Sea Drilling 
Project site 524 (Gombos, 1984). 
Paleocene - lower Eocene. Barents Sea, 100-500 m depth, between Franz- Josefs Land and Novaya 

Zembla (Grunow, 1884). (Date uncertain, more probably Paleocene.) 
Lower Eocene. Jutland, Denmark (BM 45388, 45391, 74094; PH coll. Schulze arr. 3192; Benda, 1972). 

Glyng0re, Jutland, Denmark (BM 30847). 

Skive, Jutland, Denmark (BM 14744, 69223; Tempere & Peragallo, 1891, 1913). 

Harre, Jutland, Denmark. Drill hole (Fenner, 1985). 

Limfjord, Jutland, Denmark (BM 9840, 13588, 45644, 46747, 54353, 54354, 54515, coll. Adams Bess. 
1154; Miller, 1969). 

Island of Mors, Denmark (BM 9856, 9857, 10411, 10413, 12927, 14297, 27798, 30863, 31535, 32718, 
32719, 38292, 44007, 44008, 44009, 44509, 54351, 54357, 60821, 60822, 60823, 60824, 60825, 60826, 
64013, 68374, 68375, coll. Adams Bess. 1788, F1096, GC3483, J587, J2952, SEM CB15.281-289; PH 
coll. Boyer S-1-18, Gen. coll. 12724; BRM Hdl/12; Heiberg, 1863; Kitton, 1870; Cleve & Moller, 1878; 
Tempere & Peragallo, 1889ft, 1907). 

Nykjobing, Island of Mors, Denmark (BM 12926, 31604, 54798, coll. Adams C514; Kitton, 1870; 
Cleve & Moller, 1878). 

Ejerslev, Island of Mors, Denmark (BM coll. Adams TS286, TS287, TS416). 

Island of Fur, Denmark (BM 10412, 14271, 14272, 27806, 35882, 45389, 53564, 60779, 64014, 68656, 
coll. Adams Bess. 358, F116, L22, SEM B28.988-991, B29.88-93; PH Gen coll. 10736; BRM Hdl/13; 
Heiberg, 1863; Kitton, 1870; Tempere & Peragallo, 1889fl, 1911). 

Henmoor, Niedersachsen, Germany (Schulz, 1927). 

Dornbusch, Hiddensee, Rostock, Germany (Schulz, 1927). 

Greifswalder Oie, Rostock, Germany (BRM 428/70; Schulz, 1927). 

London clay, near base (locality not indicated), England (Shrubsole & Kitton, 1881). 

Prionersk, Kaliningrad oblast, U.S.S.R. (Strel'nikova, Kaplan & Travina, 1978). 

Middle Volga basin, U.S.S.R. (Glezer et al., 1974). 

Sengilei, Ulyanovsk oblast, U.S.S.R. (Glezer, 1966). 

Region of Atlym, Berezovo, Kazym (i.e. 62°-64° N., 65°-67° E.), Tyumen'sk oblast, U.S.S.R. 
(Rudkevich, Rubina & Permyakov, 1957; Galerkina, 1959). 

North-west of Fergana, Uzbek S.S.R., U.S.S.R. (Glezer et al., 1974). 
Middle Eocene. Region of Atlym, Berezovo, Kazym, Tyumen'sk oblast, U.S.S.R. (Rudkevich, Rubina & 
Permyakov, 1957; Galerkina, 1959). 

Junction of rivers Ob and Chemashevskoi, Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Plain between the rivers Ob and Pur, Tyumen'sk oblast, U.S.S.R. (Paramonova, 1964). 

Tazovsk Peninsula, Tyumen'sk oblast, U.S.S.R (Paramonova, 1964). 



274 R. ROSS AND P. A. SIMS 

Middle-upper Eocene. Norwegian Sea, North Atlantic Ocean, 67° 47-11' N., 5° 23-26' E., 1297 m depth. 

Deep Sea Drilling Project site 338 (Dzinoridze et al., 1978). 
Upper Eocene. Norwegian Sea, North Atlantic Ocean, 67° 12-47' N. , 6° 18-38' E. , 1217 m depth. Deep Sea 
Drilling Project site 340 (Dzinoridze et al., 1978). 

Chelyabinsk oblast, U.S.S.R. (Proshkina-Lavrenko et al., 19496). 

Sverdlovsk oblast, U.S.S.R. (Proshkina-Lavrenko et al., 1949a, b). 

Ivdel area, well 130, Sverdlovsk oblast, U.S.S.R. (Glezer, 1966). 

Serov area, wells 762, 771, Sverdlovsk oblast, U.S.S.R. (Glezer, 1966). 

Koptelovo area, well 255, Sverdlovsk oblast, U.S.S.R. (Glezer, 1966). 

Luchinkin, Sverdlovsk oblast, U.S.S.R. (Vozzhennikova, 1960). 

'Kamichev', presumed to be Kamyshlov, Sverdlovsk oblast, U.S.S.R. (BM 35045, 65831, 65832, 
78364, coll. Adams TS743, TS748, TS749, SEM B27.529-530, CB13.717-725, CB15.276-280, 
CB 15. 368-375, 4106-4116; Cheneviere, 1934). 

Kimkya, basin of the river Severnaya Sos'va, Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Tyumen', Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Lar'yak, Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Region of Atlym, Berezovo, Kazym, Tyumen'sk oblast, U.S.S.R. (Rudkevich, Rubina & 
Permyakov, 1957; Galerkina, 1959). 

Junction of rivers Ob and Chemashevskoi, Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Nyda, Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Plain between the rivers Ob and Pur, Tyumen'sk oblast, U.S.S.R. (Paramonova, 1964). 

Basin of the Arka-Tab-Yakha, Tyumen'sk oblast, U.S.S.R. (Glezer, 1966). 

Tazovsk Peninsula, Tyumen'sk oblast, U.S.S.R. (Paramonova, 1964). 

Pudino, Tomsk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Vasyugan, Tomsk oblast, U.S.S.R. (Glezer, 1966). 

Parabel', Tomsk oblast, U.S.S.R. (Glezer, 1966). 

River Shet-Irgiz, bore no. 6, Kazakhstan, U.S.S.R. (Glezer, 1969). 

Region of Ayak-Kuduk, Kazakhstan, U.S.S.R. (Shibkova, 1968). , 

Region of KaraTau, Kazakhstan, U.S.S.R. (Shibkova, 1968). ^ 

Region of Lake Balkash, Kazakhstan, U.S.S.R. (Shibkova, 1968). 
Upper Eocene-Oligocene. Tyumen', Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 

Lar'yak, Tyumen'sk oblast, U.S.S.R. (Vozzhennikova, 1960). 
Lower Oligocene. Eastern slopes of the northern and central Urals - Tyumen'sk and Sverdlovsk oblasts. 
(Proshkina-Lavrenko et al., 1949a). 

Northern Urals, Tyumen'sk oblast, U.S.S.R. (BM SEM CB15.264-265, CB15.273-275). 

forma pentagona Jouse* (Figs 4, 87) 

Solium exsculptum ['exculptum'] var. pentagonatis Kitton ex Walker & Chase, Notes on some new and rare 
diatoms [I]: 5, pi. 1 fig. 10 (1886). 

Valves 5-polar. 

Paleocene. Cape Basin, South Atlantic Ocean, 29° 29-055' S., 3° 30-74' E., 4805 m depth. Deep Sea 

Drilling Project site 524 (Gombos , 1984) . 
Lower Eocene. Isle of Mors, Denmark (BM coll. Adams F1096). 

Isle of Fur, Denmark (BM 10412; Kitton, 1870; Walker & Chase, 1886). 
Upper Eocene. 'Kamichev', presumed to be Kamyshlov, Sverdlovsk oblast, U.S.S.R. (BM 65833, SEM 
CB16.491-495, CB16.789-790). 
Plain between the rivers Ob and Pur, Tyumen'sk oblast, U.S.S.R. (Paramonova, 1964). 
Tazovsk Peninsula, Tyumen'sk oblast, U.S.S.R. (Paramonova, 1964). 
Lower Oligocene, Northern Urals, Tyumen'sk oblast, U.S.S.R. (BM SEM CB15. 260-263, CB15.271- 
272). 

Although both H. L. Smith (1872) and Van Heurck (1896) included Solium in Hemiaulus, 
neither of them pubhshed the combination Hemiaulus exsculptus; this was first done by Schiitt 

* We have not been able to trace the valid publication of this name, which is used by Paramonova (1964) but assume 
that it has been given valid publication in some work not available to us. If not, it is va-idly published here by the 
reference given to a validly published varietal name. In this connexion, it should be pointed out that epithets do not have 
priority outside their own rank. 



GENERA OF THE BIDDULPHIACEAE 275 

(1896), whose publication appeared later in the year than Van Heurck's. The specific epithet has 
frequently been misspelled 'exculptum' or, in combination with Trinacria, 'exculpta' . 

The 5-polar form of this species occurs in a number of localities along with the quadripolar 
form, although normally less commonly. It has never been recorded in the absence of the 
quadripolar form. As it does not seem to have a distinct distribution in either space or time, there 
is no case for separating it taxonomically at a higher rank than forma, but we have recognized it 
at this level as some authors may wish to note that it is present in material on which they are 
reporting. The size range of the specimens of this form that we have seen corresponds to the 
lower part only of the size range of the quadripolar specimens. 

Walker & Chase (1886) attribute the name Solium exsculptum var. pentagonalis to Kitton, 
saying 'the variety with five angles is mentioned by Kitton', but he did not publish the name; he 
merely remarked: 'in the Fuur deposit valves with five angles are not uncommon' (Kitton, 1870) . 

This species shows little variation in any other characters than the number of poles except that 
specimens with prominent hollow spines or occluded processes occur only in certain localities. 
These structures are not present on specimens from the lower Eocene of Denmark but are 
usually to be found on those from 'Kamichev' (presumed to be Upper Eocene from Kamyshlov, 
Sverdlovsk oblast, U.S.S.R.). A specimen from material received from the late Mr A. L. 
Brigger and labelled 'North Urals, U.S.S.R. OHgocene' has what appears to be the base of one 
such spine or occluded process, its upper part, like the external part of the labiate process, 
having been broken off. The broken base of such a structure may be present on one of the 
specimens of the tropical Indian Ocean illustrated by Jouse (1977: tabl. 67 fig. 10) and one or two 
are present on the upper Eocene specimen from the western Siberian lowlands illustrated by 
Glezer et al. (1974: tabl. 23 fig. 4). None, however, are visible on the illustrations of specimens 
from the late Paleocene of Cape basin. South Atlantic (Gombos, 1984: pi. 8 figs 9-11), nor on 
the upper Eocene specimen from NW Siberia illustrated by Paramonova (1964: tabl. X fig. 2). 

The end valves (Figs 1, 2), i.e. those not inseparably linked to their sibhng valves by 
interlocking linking spines, differ from the others not only in their linking spines but also in the 
inclination of the external part of their labiate processes. On the valves within the chain these are 
so strongly inclined that they almost or quite touch the surface of the projection distal to their 
point of insertion (Fig. 3) ; this makes them very difficult to detect when specimens in girdle view 
are examined with the fight microscope. On end valves, however, the external tubes of the 
labiate processes are inclined at less than 45 degrees to the pervalvar axis, and they may be 
almost vertical (Fig. 1). The frequency with which end valves are encountered is such as to 
suggest that the chains in this species were short. 

We have seen two lower Eocene specimens from Glyng0re, Jutland, Denmark, that have the 
appearance of being initial valves. These have sides of 91 fxm and 94 /xm. The first has no costae 
delimiting three of its four projections whilst the other is somewhat asymmetrical, two adjacent 
projections being larger than the other two. Both of these specimens have the labiate process 
opposite the centre of a side of the valve instead of opposite a projection, the normal position. 

The specimens illustrated by A. Schmidt (1890: taf. 152 figs 24-25), of which he says 'nach 
Witt's Ansicht vielleicht zu Solium exsculptum zu rechnen' are not this species but Trinacria 
cristata Gombos (1982). We have examined the specimens that are the originals of these figures 
(BM coll. Adams J3016- both specimens are mounted on this slide). Proshkina-Lavrenko et al. 
(1949^) treat Trinacria semseyi Pant, as a synonym of Solium exsculptum but this is a manifest 
error. T. semseyi, which comes from the upper Eocene of Kuznetsk, Penza oblast, U.S.S.R. , is a 
tripolar species which has the characters of Trinacria, not Solium. 

The earliest records of this species are from the upper Paleocene of the South Atlantic 
(Gombos, 1984) and the tropical part of the Indian Ocean (Mukhina 1974, 1976). All the 
subsequent records are from northern Europe or adjacent seas, north-western Siberia, the 
central Volga basin, and Kazakhstan. It persisted throughout the Eocene in this area, and 
continued into the lower Oligocene in north-western Siberia. It is not rare in its lower Eocene 
localities nor in the upper Eocene from north-western Siberia, but elsewhere it is apparently 
infrequent in its occurrence. This rather pecuHar distribution, in low latitudes in the Paleocene 
and in high northern latitudes from the lower Eocene to the Oligocene, seems more likely to be a 



276 R. ROSS AND P. A. SIMS 

reflection of the inadequacy of the fossil record than a complete record of its history. It is 
presumably because it is only in north-western Siberia that it has been found to occur commonly 
in the upper Eocene that Fenner (1985) says that it is characteristic for the late Paleocene and 
earliest Eocene, whereas Glezer (in Glezer et al., 1974: 140) remarks that it did not decrease in 
the upper Eocene. 

A misprint in a previous paper (Ross, Sims & Hasle, 1977) that we have only recently detected 
suggests that Solium exsculptum occurs in Eocene samples from the Falkland Plateau taken by 
R/V Vema. It does not. The first line of the upper of the two paragraphs on p. 180 of that paper 
which begins: 'Hemiaulus februatus and Trinacria exsculpta . . .' should read: 'Hemiaulus 
centralitenuis, H. reflexispinosus , and H. prolongatus from . . .' 

We have seen three specimens of this species labelled as coming from localities that we have 
not included when listing its distribution. These are: PH Gen. coll. 89190, Jackson's Paddock, 
Oamaru, New Zealand; BM coll. Adams TS478, San Redondo, California, U.S.A. BRM 
Hd2/27, Santa Monica, California, U.S.A. As there is no other record of the occurrence of this 
species in any of these much studied deposits, we regard these specimens as strays and have not 
accepted these records. 

II. PSEUDORUTILARIA (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni, Syll. 

Alg. 2: 854 (1894). 

Subgen. Pseudo-rutilaria Grove & Sturt in /. Quekett microsc. Club II, 2: 324 (1886), nom invalid (Arts 

21.1,33.1). 
Rutilaria subgen. Pseudorutilaria Grove & Sturt ex De Toni & Levi in Notarisia 2: 349 (1887). 

Frustules rectangular, united in inseparable chains. Valves bipolar or tripolar, the bipolar valves 
having a straight or curved apical axis, with long projections each cut off from the central portion 
by a transverse sulcus or internal costa and crossed by a number of other sulci or internal costae; 
valve surface undulate. Elevations at each pole not very tall, not expanded above, with a clearly 
defined subocellus on their distal side, interlocking linking spines at their tip on the proximal 
side, and anastomosing low external costae on their lower part and on the mantle below them. 
Mantle with a deep furrow above a narrow hyaline band, the furrow usually continuous around 
the whole valve but sometimes present only where the valve margin is convex. Areolae 
irregularly arranged on the valve surface and the upper part of the mantle, each with a simple 
volate velum. A well-developed marginal ridge, normally bearing many strong marginal spines, 
these sometimes fused with those of the sibling valve. An irregular ring of occluded processes on 
the central portion of the valve more or less alternating with troughs with raised margins that 
clasp the inclined upper parts of the occluded processes of the sibling valve usually present but 
absent on end valves and often on the shortest valves. One or two labiate processes normally 
present on the central portion of the valve, more or less on the transapical axis in bipolar species, 
their internal openings straight or slightly curved and sessile, their external tubes stout and very 
similar to the occluded processes but reaching to the margin of the sibling valve. 

Type species: Pseudorutilaria monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni (loc. 
cit.). 

The first account of this genus was given by Grove & Sturt (1886). This was headed 'Pseudo- 
rutilaria. Nov. subgenus', but there was no positive indication of the genus to which it was to be 
attributed. The statement: 'We have formed this subgenus for the reception of the species 
described below, which, while resembling Rutilaria in other characteristics, is without the 
central clasping process, so important a feature of that genus', and the fact that the account of it 
immediately follows that of Rutilaria, make it plain that they intended it as a subgenus of that 
genus. However, the International Code of Botanical Nomenclature provides that 'the name of a 
subdivision of a genus is a combination of a generic name and a subdivisional epithet connected 
by a term . . . denoting its rank' (Art. 21.1) and that 'a combination (autonyms excepted) is not 
vahdly pubUshed unless the author definitely indicates that the epithet or epithets concerned are 
to be used in that particular combination' (Art. 33.1). Grove & Sturt did not give such an 



GENERA OF THE BIDDULPHIACEAE 277 

indication. They used the name 'Ps.-rutilaria monile' in the text for the one species that they 
included in this subgenus, and ' Pseudo-rutilaria monile' in the legend of the plate on which it is 
illustrated. 

The subgeneric name Rutilaria subgenus Pseudorutilaria was validly published by De Toni & 
Levi in 1887, as also was the specific name Rutilaria monile. They attributed both to Grove & 
Sturt and gave a Latin translation of Grove & Sturt's account of the two taxa. De Toni (1894) 
later treated Pseudorutilaria as a genus, attributing this name also to Grove & Sturt. At the same 
time he validly published the combination Pseudorutilaria monile for the type species. Grunow 
(1887), Tempere & Peragallo (1890), and Schmidt (1893) used the name Pseudorutilaria monile 
earher than this but did not give the generic name vaUd publication. 

The most striking and unusual feature of Pseudorutilaria is the ring of occluded processes and 
troughs with raised margins that clasp the distal ends of the occluded processes of the sibling 
valve. This feature is well shown in Fig. 18. The external tubes of the labiate processes resemble 
the occluded processes and are clasped by the raised margins of the troughs in the same way; 
they, however, reach the margin of the valve (see Fig. 32), whereas the occluded processes stop 
short of it. 

There are many points of similarity between this genus and Solium Heiberg: the subocelli of 
the two genera resemble each other closely; both have interlocking linking spines on the 
proximal sides of the elevations and low anastomosing external costae on the elevations and the 
upper part of the mantle, although these costae are less well developed in Pseudorutilaria than in 
Solium; in both there is a concavity of the mantle above a narrow hyaline rim; both have 
projections cut off from the central portion of the valve by sulci or internal costae, although in 
Pseudorutilaria the projections are crossed by additional transverse sulci or internal costae; both 
normally have one labiate process inserted towards the margin of the central portion of the 
valve, with an elongate sessile internal opening and a stout external tube. The two genera are 
clearly closely related. 

In some species oi Pseudorutilaria, e.g. P. incompleta R. Ross & P. A. Sims, the projections 
are cut off from the central portion of the valve and are crossed by sulci, inward folds of the valve 
(Figs 39-41). In others, e.g. P. monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De 
Toni, all the transverse structures are soUd internal costae with their free edges laterally 
expanded and level with the margins of the valve, which itself has an inwardly projecting rim 
(Figs 11, 12). In P. hendeyi R. Ross & P. A. Sims, however, the proximal transverse structures 
are sulci and the more distal ones are costae (Figs 48-50, 54). 

The only species until now added to the genus since its original publication is Pseudorutilaria 
monomembranacea Schrader in Schrader & Fenner (1976: 994). This species, however, has 
neither terminal elevations with subocelli and interlocking linking spines nor occluded processes 
and associated clasping troughs. Strel'nikova (in Dzinoridze et al., 1979: 63) erected for this 
species the new genus Praecymatosira, a genus which she considered was related to Cymatosira 
Grun., and this seems to be its correct position. 

In this paper we describe five new species of Pseudorutilaria from Eocene material from the 
south-western Atlantic. One of these is the species misidentified by Hajos (1976: 829) as P. 
monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni. Also, the diatom 
misidentified as Eunotogramma weissei var. productum Grove & Sturt by Fenner (1977: 519) is 
another of the species described here as new. 

The genus has so far been found only in middle to upper Eocene, or possibly lowest 
Oligocene, deposits in the south-western Atlantic, the Tasman Sea, and New Zealand. This 
suggests that its species may well prove to be useful stratigraphic markers. 

Key to the species 

la. Valves tripolar 6. hendeyi(p. 283) 

lb. Valves bipolar: 

2a. Central portion pentagonal and with a crenate margin 5. incompleta (p. 282) 

2b. Central portion circular and with an entire margin: 
3a. Margins of the projections crenate or undulate: 



278 R. ROSS AND P. A. SIMS 

4a . Margins of the projections crenate , indented opposite every transverse costa 

1 . moni7e(p. 278) 

4b. Margins of the projections undulate, undulations more widely spaced than the 

transverse costae 4. hannai{p. 281) 

3b. Margins of the projections with one gibbosity, elsewhere entire: 

5a. Gibbosity of the margins of the projections adjacent to the central portion 

2. clavata {p. 219) 

5b. Gibbosity of the margins of the projections about midway between the central 

portion and the apex 3. nodosa (p. 280) 

1. Pseudorutilaria monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni, Syll. 
A/g. 2:854(1894). 
(P1.2,P1. 12fig. 89) 

'Pseudo-rutilaria' monile Grove & Sturt in /. Quekett microsc. Club II, 2: 324, pi. 18 figs 7-8 (1886), 

nom. invalid (Arts 21.1, 33.1). 
Rutilaria monile Grove & Sturt ex De Toni & Levi in Notarisia 2: 349 (1887). 

Valve bipolar, straight, 75-190 /xm long, 17-24 /am broad; central portion circular, separated 
from the projections by transverse costae; projections about half the breadth of the central 
portion at their proximal ends, tapering very slightly towards the apices, margins crenulate, 
indented opposite each transapical costa, costae 6-7 fx m apart, laterally expanded at their free 
edges, which are level with the inwardly projecting valve margin. Height to the top of the 
elevations 10-14 /xm. Linking spines 3-4 on each elevation, extending rather more than 
half-way around the summits, c. 3 /xm tall. Areolae poroid, with a slightly raised external rim, 
diameter c. 0-3 /xm, scattered over the whole valve surface and the upper part of the mantle, 
sparser on the central portion than on the projections, where they are c. 8 in 10 /xm. Ring of c. 6 
occluded processes on the central portion, absent on end valves. A stout, flattened marginal 
spine with a serrate tip between each pair of marginal costae , the spines of sibling valves opposite 
and abutting against one another, but absent on end valves. Labiate process one, situated in the 
ring of occluded processes on the central portion, more or less on the transapical axis, the labiate 
processes of sibling pairs of valves on the same side. 

Middle Eocene. South-western Atlantic, Falkland Plateau, 49° 52-194' S., 40° 50-713' W., 1844 m depth. 
Deep Sea Drilhng Project site 512 (Gombos, 1983). 

Upper Eocene. Oamaru, Otago Province, New Zealand (BM 9241, 9242, 11094, 11140, 11164, 11243, 
33316 (Jackson's Paddock), 33317 (Jackson's Paddock), 46542, 46577, 46606, 46633, 52709, 52759, 
52760, 60842, 63397, 63398, 64687, 65090, 66566, 71539 (lectotype), 74150, 74151, 76748, 77754 
(Jackson's Paddock), 77756, 77758 (Allen's Farm), 77760, coll. Adams Bess. 979, C499, G80, G97 
(Cormack's Siding), G104 (Cormack's Siding), G624, GC2614, GC3208, J548, J818 (Jackson's Pad- 
dock), TS23, TS271, TS292, TS433 (Allen's Farm), TS949 (Bain's Farm, lower), SEM B4.414-415 
(Division Hill), B4.425-428 (Division Hill), CB14.200-208, 44519-44537; BRM Htl/1, Htl/2, Htl/3 
(Jackson's Paddock); Grove & Sturt, 1886; Schmidt, 1893; Tempere & Peragallo, 1890 (Bain's Farm, 
Totara); Tempere & Peragallo, 1908 (Bain's Farm); Laporte & Lefebure, 1929; Tsumura, 1964 (Flume 
Creek). 

South-western Atlantic, 49° 48-67' S, 36° 39-53' W, 5095 m depth. Deep Sea Drilling Project site 328 
(Gombos, 1976). 

Upper Eocene-lower OHgocene. South-western Atlantic, Falkland Plateau, 51° 00-28' S, 46° 58-30' W., 
2589 m depth. Deep Sea Drilhng Project site 511 (Gombos & Ciesielski, 1983). 

Although this species was described and figured by Grove & Sturt in 1886, the name Pseudo- 
rutilaria monile that they used for it was not validly published as they did not give valid 
publication to Pseudo-rutilaria as a generic name. As explained above (p. 276), they said that 
Pseudo-rutilaria was a subgenus but nevertheless made the combination with that name and not 
with Rutilaria Grev., which, as they impHed but did not definitely state, was the genus of which 
Pseudo-rutilaria was a subgenus. It was De Toni & Levi (1887) who first gave valid publication to 
a name for the species. It is clear from their protologue that it was based entirely on Grove & 
Sturt's account and its type material is thus the specimens that those authors had studied. 



GENERA OF THE BIDDULPHIACEAE 279 

Although both Grove's and Sturt's collections are in the British Museum (Natural History), 
none of the specimens in either of them is the original of either of their figures. One cannot say 
which, if any, were in their hands at the time when their paper was written; some certainly were 
not. There are a number of other slides mounted by Grove elsewhere in the collections at the 
Museum, and one of these, BM 71539 from the collection of Thomas Saxton, is a strewn slide 
from Oamaru labelled in Grove's hand. Amongst the names of species on the label is 'Rut. 
monile', and it can therefore be regarded as certain that this slide was prepared and labelled 
before Grove & Sturt published their account of the species and that the specimens on it are part 
of the original material; on all the other slides labelled by Grove the name is given as 
Pseudo-rutilaria monile. We therefore designate this slide as lectotype. 

We have seen under the scanning electron microscope one specimen of this species that we 
interpret as an end valve. This specimen, which is 120 />tm long, has no occluded processes nor 
troughs on its central portion and it lacks marginal spines. 

This species has a known range from the middle Eocene-Oligocene boundary in deposits 
from the south-western Atlantic and it also occurs in the late upper Eocene deposit at Oamaru, 
Otago Province , New Zealand , from which it was originally described . The records of it from the 
upper Eocene of the Tasman Sea by Hajos (1976) are based on a misidentification; the species 
she illustrated is Pseudorutilaria clavata R. Ross & P. A. Sims. 

2. Pseudorutilaria clavata R. Ross & P. A. Sims, sp. nov. 
(PI. 3; PI. 12 figs 90, 91) 

Pseudorutilaria monile sensu Hajos in Init. Rep. Deep Sea Drilling Proj. 35: 829, pi. 14 figs 10-12 
(1976), non Grove & Sturt. 

Valva bipolaris, recta, 65-155 /urn longa, 22-30 /Am lata; portio centralis circularis, ab projecturis sulcis 
transversis separata; projecturae sulcis transitae, utraque margine inter sulcum juxta portionem centralem 
et proximum gibboso, apices versus primo gradatim angustatatum in apices clavatos rotundatos expansa, 
margine integro. Duo sulci magis proximales in utraque projectura 10-12 /xm distantes, alii 6-8 /am 
distantes. Altitudo ad vertices elevationum 12-14 /xm. Spinae ligantes in utraque elevatione 4-5, circum 
pauUo plus quam dimidium proximalem verticis positae, supra valde expansa, 2-0-3-5 jxva altae. Areolae 
poroides, volis occlusae, extrinsecus margine elevato, diametro c. 0-3 /am, in fronte et parte superiori limbi 
irregulariter dispersae, 6-8 in 10 /u,m sed in portione central! distantiores, in latere proximali elevationum 
carentes vel rarissimae. Areolae subocellorum 40-50 in 10 /xm. Annulus ex c. 6 processibus occlusis 
constans in portione central!, in valvis curtissimis carens. Spinae marginales acutae, eis valvae continguae 
non oppositae, c. 5 in 10 yam. Rimoportula una, in annulo processuum occlusorum et plus minusve in axe 
transapicali posita, eae valvarum contiguarum idem latus versus extensae. Taenia sola visa (?valvocopula) 
indivisa, c. 8 /im alta, seriebus verticalibus areolarum parvarum c. 20 in 10 jxm, areolis c. 15 in 10 /am. 

Typus: BM 81136, ex stratis eocaenicis de profundis maris Atlantici australis. 

Valve bipolar, straight, 65-155 /am long, 22-30 /u,m broad; central portion circular, separated 
from the projections by transverse sulci; projections crossed by sulci, gibbous between the 
central portion and the next transverse sulcus, distal to this tapering and then expanded to 
clavate rounded apices, their margins entire. Two proximal transverse sulci on each projection 
10-12 /Ltm apart, the others 6-8 />tm apart. Height to the top of the elevations 12-14 /am. Linking 
spines on each elevation 4-5, extending rather more than half-way round the proximal side of 
the summit of the elevation, strongly expanded above, 2 -0-3 -5 /xm tall. Areolae poroid, 
occluded by volae, with a slightly raised external rim, diameter c. 0-3 /xm, scattered over the 
valve surface and the upper part of the mantle, 6-8 in 10 /u,m, rather sparser on the central 
portion, absent or very sparse on the proximal side of the elevations. Areolae of the subocellus 
40-50 in 10 /am. Ring of about 6 occluded processes on the central portion, absent in the shortest 
valves. Marginal spines pointed, not opposite those of the sibling valve, c. 5 in 10 /am. Labiate 
process one, situated in the ring of occluded processes and more or less in the transapical 
axis, the labiate processes of sibling valves on the same side. The only girdle band seen 
(? valvocopula) undivided, c. 8 /am deep, with vertical rows of areolae, rows c. 20 in 10 /am, 
areolae c. 15 in 10 /am. 



280 R. ROSS AND P. A. SIMS 

Middle Eocene. South-western Atlantic, Falkland Plateau, 51° 08' S., 54° 22' W., 1525 m depth. Vema 
cruise 17, core 107, (BM SEM CB12.623-628, CB13. 172-173, CB13.313-326, 5861-5878), 50 cm (BM 
81137, SEM CB14.098-101, CB14.121-124, CB14.930-931, 44263-44268, 44631-44638), 175 cm (BM 
81136). 

Middle-upper Eocene. South-western Atlantic, Falkland Plateau, 53° 01' S., 52° 52' W., 2880 m depth. 
Vema cruise 18, core 104, 330 cm (BM SEM CB13.585-594, 5891-5894, 5901-5909). 

Upper Eocene. South-western Pacific, South Tasman Rise, 47° 59-84°S, 147° 45-85' E., 1600 m depth. 
Deep Sea Drilling Project site 281 (Hajos, 1976). 

South-western Pacific, Tasman Sea, 43° 54-60' S., 154° 16-96' E., 4750 m depth. Deep Sea Drilling 
Project site 283 (Haj6s, 1976). 

This species is easily distinguished from Pseudorutilaria monile by the outline of the valve; the 
margin is gibbous between the proximal sulcus and the next and distal to this it is entire, not 
crenulate. These characters are well shown on Hajos's figures (1976: pi. 14 figs 10-12), making it 
obvious that the specimens from the upper Eocene of the south-western Pacific that she 
identified as P. monile are actually P. clavata. 

The girdle band, probably a valvocopula, shown on Fig. 19, is the only one that we have seen 
on any member of the genus. It would seem that this band is uninterrupted; all the ends of the 
three pieces of it appear to be breaks and it would seem that no part is missing. 

We have seen two specimens of Pseudorutilaria clavata that are about 65 /utm long. Both of 
these consist of two sibling valves inseparably joined by interlocking linking spines on their 
elevations (Fig. 20). There is one labiate process but no occluded processes on the central 
portion of each valve of these specimens. It would seem that in this species, as in P. incompleta 
R. Ross & P. A. Sims (see p. 282 below), these structures are not developed on the shortest 
specimens. There are, however, indications of a ring of tooth-hke spines at the centre of the 
valve similar to those on the end valves of P. incompleta, but the position of the specimens does 
not allow certain observation of this. 

Pseudorutilaria clavata has a range from the middle Eocene to the upper Eocene. It has been 
found in samples from the Falkland Plateau that date from the middle Eocene and from the 
middle-upper Eocene boundary, and in others from the Tasman Sea area of the south-western 
Pacific that are upper Eocene in date. 

3. Pseudorutilaria nodosa R. Ross & P. A. Sims, sp. nov. 
(P1.4;P1. 12fig. 92) 

Valva bipolaris, recta curvatave, 145-195 /xm longa, 32-34 ju,m lata; portio centralis circularis, ab 
projecturis costis internis transversis separata; projecturae costis transitae, apices pauUo clavatos rotunda- 
tos versus gradatim angustatae, sed medio gibbosae, marginibus alibi integris. Costae 7-10 /xm distantes, 
eae medio propriores distantiores. Altitudo ad vertices elevationum c. 15 fxm. Spinae ligantes 2-4 in 
utraque elevatione, circum dimidium proximalem verticis positae, c. 4 /tm altae. Areolae poroides, 
extrinsecus margine paullo elevato, diametro c. 0-3 fxm, in fronte et parte superiori limbi irregulariter 
dispersae, c. 6 in 10 /xm. Areolae subocellorum 40-45 in 10 fjum. Annulus ex 6-7 processibus occlusis 
constans in portione centrali. Spinae marginales acutae, eis valvae continguae non oppositae, 5-7 /xm 
distantes, sed prope apices crebriores. Rimoportula una, in annulo processum occlusorum et plus minusve 
in axe transapicali posita, eae valvarum contiguarum idem latus versus extensae. 

Typus: BM 81138, ex stratis eocaenicis de profundis maris Atlantici australis. 

Valve bipolar, straight or curved, 145-195 fxm long, 32-34 /xm broad; central portion circular, 
separated from the projections by transverse internal costae; projections crossed by costae, 
gradually tapering but gibbous about half-way to the slightly clavate rounded apices, margins 
entire. Transverse costae 7-10 /Ltm apart, those near the apices the more closely spaced. Height 
to the top of the elevations c. 15 /xm. Linking spines on each elevation 2-4, extending half-way 
around the summit on its proximal side, c. 4 fim tall. Areolae poroid, with a slightly raised 
external rim, diameter c. 0-3 /xm, scattered over the valve face and the upper part of the mantle, 
c. 6 in 10 /xm. Areolae of the subocellus 40-45 in 10 /xm. Ring of 6-7 occluded processes on the 
central portion. Marginal spines pointed, not opposite those of the sibling valve, 5-7 /xm apart 
but closer near the apices. Labiate process one, situated in the ring of occluded processes on the 



GENERA OF THE BIDDULPHIACEAE 281 

central portion and more or less in the transapical axis, the labiate processes of sibling valves on 
the same side. 

Middle Eocene. South-western Atlantic, Falkland Plateau, 51° 08' S, 54° 22' W, 1525 m depth. Vema 

cruise 17, core 107, 50 cm (BM SEM CB15.059-071), 175 cm (BM 81138). 
Middle-upper Eocene. South-western Atlantic, Falkland Plateau, 53° 01' S, 52° 52' W, 2880 m depth. 

Vema cruise 18, core 104, 330 cm (BM SEM CBll. 813-814, 5879-5883). 

? 'Lamont E8'. (Locality and age uncertain, see p. 271) (BM SEM 2715-2728). 

The swelling about half-way along each projection is the distinguishing feature of this species, 
and it is to this that the specific epithet applies. Pseudorutilaria clavata is the only other species of 
the genus in which the margins of the projections are not undulate or crenate. It, too, has a 
swelhng on the projections, but that is situated immediately adjacent to the central portion. In 
P. clavata, also, the transverse structures are sulci, not the costae that are found in P. nodosa 
(Figs 24, 26) . None of the specimens of this species that we have seen has any trace of a velum in 
its areolae. 

The certain records of this species are from samples from the Falkland Plateau that date from 
the middle Eocene and from the middle-upper Eocene boundary. We have also seen specimens 
from the sample labelled 'Lamont E8', which we believe to come from the same area and to be of 
about the same age. 

4. Pseudorutilaria hannai R. Ross & P. A. Sims, sp. nov. 
(PI. 5; PI. 12 fig. 93) 

Valva bipolaris, curvata, 150 yam longa, 32 ju-m lata; portio centralis circularis, ab projecturis costis 
transversis separata; projecturae costis transitae, ad fines suas proximales latitudine 2/3 ejus portionis 
centralis, apices angustos rotundatos versus decrescentes, marginibus undulatis, non crenulatis, undis 
15-18 ixm distantibus. Costae c. 8 /am distantes. Altitudo ad vertices elevationum c. 13-5 /u,m. Spinae 
ligantes 2-3 in utraque elevatione, circum dimidium proximale verticis positae, supra valde expansae, c. 4 
/am altae. Areolae poroides, extrinsecus margine pauUo elevato, diametro c. 0-5 /am, in f route et parte 
superiori limbi irregulariter dispersae, 6-8 in 10 /xm, in portione centrali distantiores. Annulus ex 3-4 
processibus occlusis constans in portione centrali. Spinae marginales 2-3 in 10 /am, pro parte maxima 
clavatae et eis valvae contiguae non oppositae et nonnullae cum eis implexae, aliquae irregulariter 
formatae positaeque. Rimoportula una, in annulo processuum occlusorum et plus minusve in axe 
transapicali, positae, in valvis nonnuUis carentes. 

Typus: BM 81300, ex stratis eocaenicis de profundis maris Atlantic! australis. 

Valve bipolar, curved, 150 /am long, 32 /am broad; central portion circular, separated from the 
projections by transverse costae; projections crossed by costae, at their proximal ends about % 
the width of the central portion, tapering to narrow rounded apices, their margins undulate, not 
crenulate, the crests of the undulations 15-18 /am apart. Costae c. 8 /am apart. Height to the top 
of the elevations c. 13-5 /am. Linking spines 2-3 on each elevation, extending half-way round the 
proximal side of the summit, strongly expanded above, c. 4 /am tall. Areolae poroid, with a 
slightly raised external rim, diameter c. 0-5 /am, scattered over the valve face and the upper part 
of the mantle, 6-8 in 10 /am, rather sparser on the central portion. Ring of 3-4 occluded 
processes on the central portion. Marginal spines mostly clavate and alternating and in places 
interlocking with those of the sibUng valve, 2-3 in 10 /am, but some irregular in shape and 
arrangement. Labiate process one, situated in the ring of occluded processes, and more or less in 
the transapical axis, absent on some valves. 

Middle-upper Eocene. South-western Atlantic, Falklands Plateau, 53° 01' S., 52° 52' W., 2880 m depth. 
Vema cruise 18, core 104, 330 cm (BM 81300, SEM CB13.595-599, CB13.601-607, 5910-5912). 

This species is the rarest of the genus and we have seen only one specimen which we have 
examined with the scanning electron microscope and then mounted to serve as the type. The 
undulate outline, which is not related to the position of the transverse costae (Fig. 34), 
distinguishes it from the other bipolar species. The one specimen that we have seen consists of 
two joined sibling valves. One of these has one labiate process, the other appears to have no 



282 R. ROSS AND P. A. SIMS 

labiate process. We have not seen a specimen of any other species of the genus without a labiate 
process. This specimen comes from a middle to late Eocene sample from the Falkland Plateau. 

5. Pseudorutilaria incompleta R. Ross & P. A. Sims, sp. nov. 
(PI. 6; PI. 13 figs 94, 95). 

Eunotogramma weissei var. productum ['producta'] sensu Fenner in Init. Rep. Deep Sea Drilling Proj. 
39: 519, pi. 27 figs 3-5 (1977), non Grove & Sturt. 

Valva bipolaris, angulo 30°-45° inter projecturas, margine ubique crenulato, 50-155 fjum longa, 25-45 /am 
lata; portio centralis pentagonalis, ab projecturis sulcis transversis sed non indentationibus marginis 
separata; projecturae sulcis transitae, apices angustos rotundatos versus gradatim angustatae. Sulci 6-7 
fim distantes. Altitude ad vertices elevationum 8-5-11 fim. Spinae ligantes in utraque elevatione 1-2, 
plerumque supra expansae et implexae, c. 2 /xm altae, aliquando spina una decrescens c. 3-5 (xm alta. 
Areolae poroides, volis occlusae, extrinsecus margine elevato, diametro c. 0-3 /u,, in f route et parte 
superior! limbi irregulariter dispersae, 6-10 in 10 /xm. Areolae subocellorum c. 50 in 10 /am. Annulus ex 
8-12 processibus occlusis constans in portione centrali, aliquando annulo spinarum humilium cuneatarum 
substitutus, in valvis curtissimis carens. Spinae marginales latae, supra expansae, eas valvae contiguae 
tangens, margine superior! irregulariter indentatae. Rimoportula una, paullo extra annulum processuum 
occlusorum et latus dorsale valvae versus posita. 

Typus: BM 81139, ex stratis eocaenicis de profundis maris Atlantici australis. 

Valves bipolar, with the projections at an angle of 30°-45° to one another, margin crenulate 
throughout, 50-155 /xm long, 25-45 /xm broad; central portion pentagonal, separated from the 
projections by transverse sulci but not by indentations of the margin; projections crossed by 
sulci, gradually tapering to the narrow rounded apices. Transverse sulci 6-7 /xm apart. Height to 
the summit of the elevations 8-5-11 /am. Linking spines 1-2 on each elevation, normally 
expanded above and interlocking, c. 2 /am tall, sometimes a tapering spine c. 3-5 /am tall. 
Areolae poroid, occluded by volae, with a raised external rim, diameter c. 0-3 /am, scattered 
over the valve surface and the upper part of the mantle, 6-10 in 10 /am. Areolae of the subocellus 
c. 50 in 10 /am. Ring of 8-12 occluded processes on the central portion, sometimes replaced by a 
ring of low cuneate spines, absent in the shortest valves. Marginal spines broad, expanded 
above, abutting against those of the sibling valve and with irregularly indented upper margins. 
Labiate process one, sUghtly outside the ring of occluded processes on its dorsal side. 

Middle Eocene. South-western Atlantic, Falkland Plateau, 51° 08' S, 54° 22' W, 1525 m depth. Vema 
cruise 17, core 107 (BM 81139, SEM B26.67-77, B26.645-651), 50 cm (BM 81140, CB14.095-097, 
CB14.102-110, CB14.125-128, CB14.909-929, CB15.041-057), 170 cm (BM 81141). 

South-western Atlantic, south-west edge of the Sao Paulo Plateau, 28° 17-22' S, 41° 05-28' W, 3203 m 
depth. Deep Sea Drilling Project, site 356 (Fenner, 1977). 
Middle-upper Eocene. South-western Adantic, Falkland Plateau, 47° 45-7' S, 57° 38-5' W, 3650 m depth. 
Conrad cruise 12, core 237 (BM SEM 5444-5450, 23545-23554, 33543-33549). 

? 'Lamont E8'. (Locality and age uncertain, see p. 271) (BM SEM 2706-2709, 2711-2714, 2729- 
2730). 

Amongst the specimens of this species that we have examined with the scanning electron 
microscope there are two which differ from the others in a number of ways: the ring of occluded 
processes and flanged troughs on the central portion is replaced by a rather smaller hyahne area 
surrounded either by eight or nine low cuneate spines or by a low flange-hke costa (Figs 39, 45); 
the external tube of the labiate process is straight, not curved; the marginal ridge is undulate but 
bears no spines; there is one spine on each elevation and this is not expanded above but tapers to 
a point. We interpret these as separation valves. A third specimen consisting of a single valve 
mounted with its inner side uppermost seems also to be a separation valve. It has a raised (seen 
as depressed from the inside) area in the centre similar in size to the raised hyaline area of the 
other two and it has no marginal spines. 

On shorter valves neither occluded processes and their associated troughs nor cuneate spines 
are present on the central portion of the valve. Of specimens we have examined with the light 
microscope, there is one that is 100 /am in length that has neither, and another of the same 



GENERA OF THE BIDDULPHIACEAE 283 

length, the holotype, which has a ring of occluded processes and troughs. A further specimen 80 
ixm long has neither occluded processes nor cuneate spines and there is no sign of any of these on 
the three specimens figured by Fenner (1977) under the name Eunotogramma weissei var. 
productum Grove & Sturt; these are 50 ^tm, 60 /xm, and 80 /xm long. E. weissei var. productum 
differs from Pseudorutilaria imcompleta in a number of ways: it has entire margins; its 
projections do not taper towards the apices and are much narrower proximally relative to the 
breadth of the central portion; they are crossed by costae expanded laterally at their free edges; 
there is a ring of c. eight labiate processes on the central portion; the elevations bear pseudocelli 
but no linking spines. Although neither linking spines nor subocelli can be seen on Fenner's 
figures, they do not show that the specimens did not possess them, and in all other respects the 
specimens that she illustrates have the characters that distinguish P. incompleta from E. weissei 
var. productum; they clearly are the former. 

Pseudorutilaria incompleta occurs in samples from the Falkland Plateau and from the 
south-west edge of the Sao Paulo Plateau that date from the middle Eocene. It also occurs in a 
sample from the edge of Falkland Plateau (Conrad cruise 12, core 237) the age of which is less 
well established but is almost certainly middle or upper Eocene. We have also seen specimens 
from 'Lamont E8', a sample for which there are no firm data but which we believe to come from 
the Falkland Plateau area and to be of about the same age as the other samples in which the 
species occurs. 

6. Pseudorutilaria hendeyi R. Ross & P. A. Sims, sp. nov. 
(P1.7;P1. 13fig. 96) 

Valva tripolaris, marginibus concavis, crenulatis, 95-185 /u,m ab apice ad apicem; portio centralis 
hexagonalis, ab projecturis sulcis transversis sed non indentationibus marginis separata; projecturae apices 
rotundatos versus angustatae, prope portionem centralem sulcis, apices versus costis, transitae. Sulci 
costaeque c. 8-5 ju-m distantes. Crenae marginis c. 11 /Am distantes, eae in lateribus oppositis projecturae 
non semper oppositae. Altitudo ad vertices elevationum c. 10 /u,m. Spinae ligantes 3-4 in utraque 
elevatione, supra expansae et implexae. Areolae poroides, diametro c. 0-3 /^m, in fronte irregulariter 
dispersae vel in seriebus obliquis indistinctis positae, 6-8 in 10 jxm, prope centrum valvae et in parte 
superior! limbi distantiores vel carentes. Area irregularis diametro c. 10 )u,m ad centrum valvae porellis c. 
30 in 10 ^tm instructa. Annulus ex c. 12 processibus occlusis constans ad centrum valvae, aream porellorum 
cingens. Spinae marginales c. 4 /am distantes, irregulariter positae, eae valvae contiguae non oppositae, 
supra dendriticales et hie illic implexae. Rimoportula una vel duae approximatae in annulo processuum 
occlusorum posita. 

Typus: BM 81142, ex stratis eocaenicis de profundis maris Atlantici australis. 

Valves tripolar, with concave crenulate margins, 95-185 )u,m from apex to apex; central portion 
hexagonal, separated from the projections by sulci but not by indentations of the margin; 
projections tapering to the rounded apices, crossed by sulci grading distally into costae. Sulci 
and costae c. 8-5 /xm apart. Crenulations of the margin c. 11 /^m apart, not always opposite on 
the two sides of a projection. Height to the top of the elevations c. 10 /xm. Linking spines 3-4 on 
each elevation, expanded above and interlocking. Areolae poroid, diameter c. 0-3 /xm, 
scattered or in indistinct oblique rows on the valve surface, 6-8 in 10 yam, sparser to absent near 
the centre of the valve and on the upper part of the mantle. An irregular area, diameter c. 10 /am, 
of porelli c. 30 in 10 /xm in the centre of the valve. Ring of c. 12 occluded processes on the central 
portion surrounding the area of porelli. Marginal spines c. 4 /xm apart, irregularly spaced, not 
opposite those of the sibUng valve, dendritically branched above and in places interlocking. 
One, or two adjacent, labiate processes in the ring of occluded processes. 

Middle Eocene. South-western Atlantic, Falkland Plateau, 51° 08' S., 54° 22' W., 1525 m depth. Vema 
cruise 17, core 107, 50 cm (BM 81143, SEM CB14.517-520), 170 cm (BM 81142). 

Middle-upper Eocene. South-western Atlantic, Falkland Plateau, 53° 01' S., 51° 52' W., 2880 m depth. 
Vema cruise 18, core 104, 330 cm (BM SEM CB. 13.576-584, CB13.617-618, 5884-5890, 5895-5900). 

This is the only species of Pseudorutilaria with an area of porelli in the centre of the valve (Fig. 
51). It would seem that the valve is thinned in this area as it is broken away in most of the 



284 R. ROSS AND P. A. SIMS 

Specimens we have seen. In none of these is there any trace of vela in the areolae and in all of 
them the subocelli have been broken away. 

Pseudorutilaria hendeyi has only been found in samples from the Falkland Plateau that date 
from the middle Eocene to the middle-upper Eocene boundary. 

III. MONILE R. Ross & P. A. Sims, gen. nov. 

Valvae bipolares, sulcis transitae, ad apices elevationibus munitae. Elevatio utraque annulo ex dentibus 
radialibus humilibus constanti circum verticem suum instructa et subocellum ferens. Limbus ad apices, 
centre valvae oppositus et in locis aliquis ubi ambitus valvae convexus est concavus, alibi verticalis. 
Areolae poroides, cribris occlusae. Rimoportulae paucae, ad centrum valvae irregulariter aggregatae. 
Taenia sola visa areolata, antiligulam ferens. 

Species typica: Monile laurentii R. Ross «fe P. A. Sims, infra. 

Valves bipolar, crossed by sulci, with elevations at the apices. Elevations with a ring of radial low 
teeth around their summits and each bearing a subocellus. Mantle concave at the apices, 
opposite the centre of the valve and in other places where the outline of the valve is convex, 
elsewhere vertical. Areolae poroid, occluded by cribra. Labiate processes few, in an irregular 
group at the centre of the valve. The only girdle band seen areolate, antiligulate. 

This genus, like Pseudorutilaria monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex 
De Toni, has something of the appearance of a row of beads. We have therefore chosen as its 
generic name the noun used in apposition as a substantival epithet for that species, a word 
meaning necklace. 

Monile resembles Pseudorutilaria in many ways but differs in important respects: its cells are 
not united in inseparable filamentous colonies, although the flattened summits of the elevations 
suggest that they did form filaments (Figs 63, 64); there are no occluded processes with their 
associated troughs at the centre of the valve (Fig. 61); there are no interlocking linking spines on 
the elevations (Figs 63, 64); also the subocelli extend on to the summits of the elevations. 
Pseudorutilaria is found earher in the fossil record than Monile, and the nature and degree of the 
relationship between the two is far from clear. It is discussed below (p. 291). 

1. Monile laurentii R. Ross & P. A. Sims, sp. nov. 
(PI. 8; PI. 13 figs 97, 98) 

Valva bipolaris, longa angustaque, marginibus sinuosis, apicibus subclavatis, ex portione centrali circulari 
tholiformi et projecturis ab portione centrali sulcis separatis, sulcis transitis et inter sulcos tholiformibus, 
constans, elevatione ad apicem utremque munita, 63-220 /xm longa, 20-25 /xm lata. Elevationes 
verticales, cylindricae, supra expansae, verticibus applanatis, utraque annulo ex dentibus humilibus 
radialibus linearibus constanti circum verticem instructa, et subocellum in vertice et in parte superiori 
expansa ferens. In utraque elevatione sub subocello costae humiles irregulariter anastomosantes, reticu- 
lum debiliter effectum facientes. Altitudo ad vertices elevationum 18-40 ^tm. Limbus ad apices, centro 
valvae oppositus et proximaliter sed non distaliter in projecturis ubi ambitus valvae convexus est, proxime 
super marginem concavus, alibi verticalis. Crista marginalis humilis, hyalina ab elevationibus ad por- 
tionem centralem extensa. Areolae poroides, cribris occlusae, extrinsecus margine paullo elevato, diam. c. 
0-7 /xm, in fronte, in elevationibus et in parte superiori limbi ubi ambitus valvae convexus est irregulariter 
dispersae, 4-6 in 10 ^tm; areolae subocellorum 15-20 in 10 jxxn, diametro c. 0-1 /u,m. Fori interstriales in 
fronte irregulariter dispersi. Spinae superficiales ad 4 /xm longae in fronte praeter partes distales 
projecturarum, ex marginibus elevatis areolarum orientes. Rimoportulae 4, ad centrum valvae irregulari- 
ter aggregatae, unaquaeque apertura interna recta paullo elevata et, extrinsecus, ex tubo crasso 4 /xm 
minimum alto et, pro parte maxima, basi fultura munito constans. Taenia sola visa c. 8 /xm alta, ad unum 
apicem antiligula valida instructa, seriebus verticalibus areolarum ornata, seriebus areolisque 10-12 in 10 
/im. 

Typus: BM coll. Adams H941, specim. 1, 12, ex depositione miocenica ad 'St. Laurent-la-Vernede', Card, 
Gallia. 

Valve bipolar, long and narrow, with sinuous margins and sub-clavate apices, consisting of a 
circular domed central portion separated by sulci from the projections that are themselves 



GENERA OF THE BIDDULPHIACEAE 285 

crossed by sulci and domed between them, and with an elevation at each apex, 63-220 /xm long, 
20-25 ^tm broad. Elevations vertical, cylindrical, expanded above, with flattened summits 
bearing a ring of low radial linear teeth around their margins, and with a subocellus on their 
summits and expanded upper part. A poorly developed network of low, irregularly anastomos- 
ing costae on the elevations. Height to the top of the elevations 18-40 /xm. Mantle strongly 
concave immediately above the margin at the apices, opposite the centre of the valve and 
proximally but not distally on the projections where the outline of the valve is convex, elsewhere 
vertical. A low hyaline marginal ridge extending from the elevations to the central portion. 
Areolae poroid, occluded by cribra, with a slightly raised external rim, diameter c. 0-7 /i,m, 
scattered over the valve surface, the elevations, and the upper part of the mantle where the valve 
outline is convex, 4-6 in 10 ixm; areolae of the suboceUi 15-20 in 10 /xm, diameter c. 0-1 fxm. 
Scattered interstrial pores on the valve surface. Superficial spines up to 4 fim long arising from 
the raised rim of the areolae on the valve surface, except on the distal parts of the projections. 
Labiate processes 4, in an irregular group in the centre of the valve, each with a straight, slightly 
raised internal opening and, externally, a stout tube at least 4 /xm tall, mostly with a single 
buttress at the base. One girdle band seen, about 8 /am deep, with a well developed antiligula at 
one apex and with vertical rows of areolae, rows and areolae 10-12 in 10 fxm. 

Miocene. St Laurent-la- Vernede, Card. France (BM 81303, coll. Adams H941, TS961, SEM CB13.634- 
650,36674-36682). 

IV. MALUINA R. Ross & P. A. Sims, gen. nov. 

Valva bipolaris, ad apices elevationibus aliquanto crassis, non supra expansis, ad «ummum utraeque 
elevationis in latere suo distali subocello minimo. Limbus verticalis. Areolae poroides, parvae, in partibus 
valvae sparsae carentesve. Spinae ligantes 2-4 in latere proximal! verticis utraeque elevationis, supra 
expansae et implexae. Rimoportula una, prope centrum valvae posita, infra sessilis longa angusta, extra 
tubo curto recto. 

Species typica: Maluina centralitenuis (R. Ross & P. A. Sims) R. Ross & P. A. Sims, comb. nov. infra. 

Valve bipolar, with moderately stout elevations at the apices, not expanded above. On the distal 
side of the top of each elevation a very small subocellus. Mantle vertical. Areolae poroid, small, 
sparse or absent on parts of the valve . Linking spines 2-4 on the proximal sides of the summits of 
each elevation, expanded above and interlocking. Labiate process single, situated near the 
centre of the valve, internally sessile, long and narrow, externally with a short, straight tube. 

We have chosen a name for this genus from the only locality where it has been found, the 
Falkland (Malvinas) Plateau. 

It is clear that the species for whicH this genus is erected here cannot be retained in Hemiaulus 
Heiberg. The interlocking linking spines and, even more obviously, the subocellus with its very 
close areolae or porelli and its definite margin are inconsistent with that position for it. Also, it 
cannot be transferred to any of the other genera with a small number of interlocking linking 
spines on each elevation. Briggera R. Ross & P. A. Sims and Dicladiopsis De Toni have 
pseudocelli with indefinite margins and expanded tips to their elevations; Solium Heiberg and 
Pseudorutilaria (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni have projections 
cut off from a central portion by sulci or costae , an indented mantle , and anastomosing costae on 
the elevations and mantle. These differences, in our view, preclude placing Hemiaulus centra- 
litenuis R. Ross & P. A. Sims in any of these genera. 

1. Maluina centralitenuis (R. Ross & P. A. Sims) R. Ross & P. A. Sims, comb. nov. 
(PI. 9) 

Hemiaulus centralitenuis R. Ross & P. A. Sims in Beih. Nova Hedwigia 54: 182, pl. 2 figs 7-9 (1977). 

Valves elliptical, 20-55 fxm long, 15-25 fxm broad, 10-12 /am tall at the domed centre, with 
more or less strongly developed acute pseudo-antiligulae at the apices and a domed centre about 
% the width of the valve in diameter. Elevations arising at the apices, cylindrical and tapering 
slightly upwards, c. 5 /am in diameter; height to the top of the elevations 15-25 /am. Valve 



286 R. ROSS AND P. A. SIMS 

surrounded by a flange-like marginal ridge. Areolae poroid, with a weak external rim, c. 0-15 
ixm in diameter, in radial rows on the central dome and sometimes also around the outer part of 
the valve face, rows 25-30 in 10 /xm, areolae 30-35 in 10 fxm; areolae also dense, 25-30 in 10 )u,m, 
on the distal side and the lower part of the proximal side of the elevations, elsewhere on the 
elevations sparse; areolae absent in a broad area around the central dome. A very small, 
transversely widened subocellus on the distal side of each elevation at its tip, the areolae or 
porelli c. 60 in 10 fim. Linking spines 2-4, interlocking, expanded immediately above their base, 
narrowed above this and then expanded into a clavate tip rounded at the apex, 6-7 /xm long. 
Labiate process about half-way between the centre of the valve and the edge of the central 
dome, slightly nearer one apex than the other. 

Middle Eocene. South-western Atlantic, Falkland Plateau, 51° 08' S., 54° 22' W., 1525 m depth. Vema 
cruise 17, core 107, 50 cm (BM SEM CB14.074-077, CB14.080-083, CB14.502-505, 5818-5826; Ross, 
Siins&Hasle, 1977). 

Middle-upper Eocene. South-western Atlantic, Falkland Plateau, 53° 01' S., 52° 52' W., 2880 m depth. 
Vema cruise 18, core 104, 330 cm (BM 78000, holotype; Ross, Sims & Hasle, 1977). 

Since we first described this diatom as a species of Hemiaulus (in Ross, Sims & Hasle, 1977), we 
have been able to examine additional, and less eroded, specimens with the scanning electron 
microscope. These have provided us with more information about the structure of the valves and 
hence we have provided a new description here. In 1977 we drew attention to the small area at 
the tips of the elevations that was broken away in the specimens that we had then seen and 
suggested that a pseudocellus might have been present there. We now know that this area is 
occupied by the structure of the sort which we have here called a 'subocellus' (Fig. 68). 

This species, and genus, is known only from cores taken from the Falkland Plateau. The 
samples in which it has been found come from the middle Eocene and the middle-upper Eocene 
boundary, 

V. BONEA R. Ross & P>. A. Sims, gen. nov. 

Frustula rectangularis, in catenis inseparabilibus conjuncta. Valvae tripolares, ad apices elevationibus 
humilibus. Elevatio unaquaeque vertice annulo perfecto spinarum ligantium implexarum circumcincto, et 
pseudocellum ferens. Limbus verticalis. Areolae poroides, rotis occlusae. Rimoportulae paucae, prope 
centrum valvae positae. Cingulum ex taeniis duis areolatis, probabiliter indivisis constans. 

Species typica: Bonea simulans R. Ross & P. A. Sims, infra. 

Frustules rectangular, united in inseparable chains. Valves tripolar, with low elevations at the 
corners. Elevations with their summits surrounded by a complete ring of interlocking linking 
spines, and bearing a pseudocellus. Mantle vertical. Areolae poroid, occluded by rotae. Labiate 
processes few, situated near the centre of the valve. Girdle bands two, areolate, probably 
undivided. 

We have named this genus after Mr E. C. P. Bone of Portslade, who has assisted us materially by 
providing us with many selected specimens of species that we have studied. 

Bonea has many points of resemblance to Strelnikovia R. Ross & P. A. Sims (1985) but differs 
in having a complete ring of interlocking linking spines on the summits of the elevations (Figs 73, 
75). These hnking spines, moreover, have bases that extend circumferentially, not radially as in 
Strelnikovia; in this, Bonea resembles more closely Briggera R. Ross & P. A. Sims (1985) and 
Dicladiopsis De Toni (1894). Keratophora Pantocsek (1889) and Thaumatonema Greville 
(1863), which, like Bonea, have a complete ring of interlocking linking spines around the 
summits of their elevations, also have the bases of these extending radially, as in Strelnikovia. 
The tall, tubular, hyaline elevations of these two genera distinguish them even more sharply 
from Bonea, which, nevertheless, is related to the group of genera that consists of them along 
with Strelnikovia, Briggera, and Dicladiopsis. 



GENERA OF THE BIDDULPHIACEAE 287 

1. Bonea simulans R. Ross & P. A. Sims, sp. nov. 
(PI. 10; PI. 13 figs 99, 100) 

Valva tripolaris, lateribus rectis vel paullo convexis, fronte ad centrum paullo tholiformi, inter centrum et 
medium uniuscuiusque lateris convexa, elevationes versus concava et ad vertices earum ascendenti; 
longitude lateris 115-125 /u,m. Elevationes sub-triangulares, interdum a limbo depressione angusta 
horizontali separatae. Altitudo ad vertices elevationum c. 30 /am. Spinae ligantes in unaquaeque 
elevatione 15-20, supra expansae et ambitu irregulari. Crista marginalis humilis, hyalina ab elevatione ad 
elevationem extensa, spinas marginales acutas, 10-12 ^im altas, c. 5 /nm distantes ferens; hae spinae 
oppositae et superimpositae sed non implexae. Areolae poroides, rotis radiis 7-10 instructis occlusae, 
radiis ramulo uno vel ramulis duobus conjunctis, diametro 0-8-1 -5 /xm, in fronte in seriebus radialibus 
dispositae, seriebus 2-5-4 in 10 /am, areolis 2-4 in 10 /am, sed prope centrum valvae distantiores; series 
areolarum media laterum versus distantiores quam eae apices versus directae; medio in valva area hyalina 
irregularis, diametro c. 10 /am; in limbo prope marginem series una horizontalis areolarum 2-5-3 in 10 /am 
et, prope cristam marginalem, alter series similaris. Fori interstriales in fronte irregulari ter dispersi. 
Pseudocellus in vertice et latere distali uniuscuiusque elevationis et in limbo sub elevatione, interdum area 
hyalina verticali in limbo et, quum depressio horizontalis inter limbum et elevationem adest, area hyalina 
super banc interruptus; areolae pseudocellorum in seriebus dispositae, seriebus in vertices elevationum 
plus regularibus quam alibi; series areolaeque 8-10 in 10 /am; areolae in vertices elevationum diametro 
0-2-0-4 /am, minor quam eae alibi diametro 0-5-1-0 /am. Rimoportulae 2-4, prope aream centralem 
hyalinam irregulariter dispositae, unaquaeque apertura interna recta, curta, paullo elevata, extrinsecus ex 
tubo verticali c. 6 /am alto appendice acuta lanceolata ad apicem instructo constans. Valvocopula parte 
exteriori c. 20 /am alta, ad omnes apices antiligula rotundata humili instructa, et areolis irregulariter 
dispersis 2-5-4 in 10 /am ornata; pleura parte exteriori c. 10 /am alta et parte interiori hyalina c. 4 /am alta, 
pars exterior serie areolarum c. 5 in 10 /am secus marginem proximalem et, huic distalibus, seriebus duabus 
paullo irregularibus areolarum c. 4 in 10 /am. 

Typus: BM 81301, ex stratis eocaenicis ad 'Carlovo, U.S.S.R.' 

Valve tripolar, with straight or slightly convex sides, the face slightly domed in the centre, 
convex between the centre and the middle of each side but concave towards the elevations and 
rising to their summits, length of side 115-125 /u,m. Elevations sub-triangular, sometimes 
separated from the mantle by a narrow horizontal depression. Height to the summit of the 
elevations c. 30 /u,m. Linking spines 15-20 on each elevation, expanded and rather irregularly 
shaped above, 2-4 /xm tall. A low hyaline marginal ridge running from elevation to elevation 
and bearing acute marginal spines 10-12 /u-m tall and c. 5 /nm apart, the spines opposite and 
overlapping but not interlocking. Areolae poroid, occluded by rotae with 7-10 spokes, the 
spokes connected by one or two side branches, diameter 0-8-1-5 /am; radial rows of areolae on 
the valve face, rows 2-5-4 in 10 /am, areolae 2-4 in 10 /am, except near the centre of the valve, 
where they are sparser; the rows directed towards the middles of the sides more widely spaced 
than those directed towards the apices; an irregular central hyaline area c. 10 /am in diarneter. 
Mantle with a horizontal row of areolae 2-5-3 in 10 /am close to the margin and another close to 
the marginal ridge. Scattered interstrial pores on the valve surface. Pseudocellus on the summit 
and distal face of the elevations and on the mantle below the elevations, where it is sometimes 
interrupted by a vertical hyaline area, a hyaline area also above the transverse furrow separating 
mantle and elevation when this is present; areolae of the pseudocelli arranged in rows, more 
regular on the summits of the elevations than elsewhere, rows and areolae 8-10 in 10 /am; 
areolae on the summits of the elevations 0-2-0-4 /am in diameter, smaller than those on the distal 
face and the mantle, which are 0-5-1-0 /am in diameter. Labiate processes 2-4, irregularly 
distributed a Uttle outside the central hyaline area, with a slightly raised, short, straight internal 
opening and, externally, a vertical tube c. 6 /am tall with a lanceolate acute appendage on its tip 
c. 2 /am long. Valvocopula with a pars exterior c. 20 /am deep, with a slightly rounded antiligula 
at each apex and irregularly scattered areolae 2-5-4 in 10 /am; pleura with a pars exterior c. 10 
/am deep and a hyaline pars interior c. 4 /am deep, the pars exterior with a row of areolae c. 5 in 10 
/am along its proximal edge and two somewhat irregular rows c. 4 in 10 /am distal to this. 

? Lower Eocene. 'Carlovo'. U.S.S.R. (BM 81301, SEM 25006-25016, 36571-36580). (For this locality see 
Ross & Sims 1985: 281) 



288 R. ROSS AND P. A. SIMS 

Bonea simulans resembles Triceratium exornatum Greville (1865), which was described from 
the Cambridge Estate, Barbados; it also occurs in samples from Springfield and Joe's River on 
that island. The material from the Cambridge Estate has been dated as middle Eocene and that 
from Joe's River as coming from the upper Eocene-OUgocene boundary (Holmes & Brigger, 
1979). There is no firm date available for the material from Springfield. We have examined the 
holotype of T. exornatum (BM 3079) and a number of other specimens of the species with the 
Ught microscope but have found no trace of linking spines or marginal spines on any of these. 
There are a few other minor differences: the areolae of T. exornatum are 4-5-5 in 10 fxm on the 
parts of the valve surface between the centre and the elevations, whereas those of B. simulans 
are 3-4 in 10 {xm on that part of the valve; there is only one row of areolae on the mantle in T. 
exornatum, but two in B. simulans (Fig. 74); in T. exornatum a transverse furrow separating 
elevation from mantle is never present; we could not detect any labiate process on T. exornatum. 
These are the only differences between the species; in outline and contour, in shape of the 
elevations, in arrangement of the areolae, and in the position and size of the pseudocelli they are 
alike. We have chosen the epithet simulans for the species described here because of this 
resemblance. It should, perhaps, be pointed out here that T. exornatum is not correctly placed in 
the genus Triceratium Ehrenberg (1839). It belongs either in Sheshukovia Glezer (1975) or in 
Biddulphia Gray (1821); we have reservations about whether these two genera are distinct. 

As we said earlier (Ross & Sims, 1985: 281), there is uncertainty about the locality from which 
the material labelled 'Carlovo' came and about its geological age. This is the only deposit in 
which we have found Bonea simulans. 

Glezer (in Glezer et al. , 1974) records Triceratium exornatum from the early to late Eocene of 
the west Siberian plain (pp. 112, 118, 119, 122, tabl. XXI fig. 1) and from the late Eocene of 
Kazakhstan (pp. 127, 129, tabl. XXXII fig. 8). The two figures are of the same specimen, that on 
tabl. XXXII rotated 120° anti-clockwise as compared with that on tabl. XXI. It is impossible to 
tell from these illustrations whether the specimen shown is T. exornatum or Bonea simulans; it 
might be either. 



VI. DEXTRADONATOR R. Ross & P. A. Sims in Bacillaria 3: 115 (1980). 

Additional information about Dextradonator eximius (Grunow) R. Ross & P. A. Sims, the type 
species of this genus, is given in the description below. 

1. Dextradonator eximius (Grunow) R. Ross & P. A. Sims in Bacillaria 3: 118, figs 1-4 (1980). 
(PI. 11) 

Syringidium eximium Grunow in Van Heurck, Syn. Diat. Belg. : pi. 106 figs 1, 3 (1883). 

Frustules cylindrical, united in inseparable chains, the apical axis 3-10 times as long as the 
diameter. Valves circular or almost so, the surface flat to concave, diameter 30-75 fxm, height to 
the centre of the valve 20-30 fim. Elevations two, marginal, cyhndrical, 5-12 /am in diameter, 
height to the top of the elevations 55-75 /Am. A hyaline marginal ridge c. 10 /xm tall and turned 
inwards along its upper edge extending from elevation to elevation. Mantle vertical, hyaline. 
Areolae poroid, c. 0-8 fim in diameter, vela eroded on all specimens seen, in radial rows on the 
valve surface, rows and areolae 9-10 in 10 /^m, extending up the proximal side of the elevations; 
a few small areolae or pores, 0-2-0-4 /xm in diameter on the distal side of the elevations near 
their summits. An irregular hyaline area containing 1-3 areolae in the centre of the valve, 
diameter 4-6 fxm. Two flattened, acute Unking spines on each elevation, partially encircling the 
elevation of the sibling valve , those on one of a pair of sibling valves arising from the distal side of 
the tops of the elevations, those on the other from the proximal side, linking spines 15-30 /xm 
tall. No labiate process seen. Cingulum consisting of two entire bands; the valvocopula 60-75 
fxm deep, its pars interior hyaline and overlapping a rounded internal ridge at the margin of the 
valve, its pars exterior with a hyaline distal margin 2-5 /xm deep, elsewhere with scattered large 
areolae c. 2 fim in diameter and 3-5 /xm apart; pleura hyaline, 15-30 ptm deep. 



GENERA OF THE BIDDULPHIACEAE 289 

Middle Eocene. Cambridge Estate, Barbados (BM 10453, 13789; Grunow in Van Heurck, 1883). 
Middle-Upper Eocene boundary. Indian Ocean, 10° 25' S, 63° 15' E, 3115 m depth, dredge sample. 

Dodo-123-Dl (BM SEM B26. 387-392, CBll. 301-302, CB19. 911-912, 46522-46532). 
Middle Eocene-lower Miocene. Barbados (BM coll. Adams TS 266; PH coll. Shulze arr. 196, 1352, 2404, 
2406, 2791). 
Bissex Hill, Barbados (BM coll. Adams TS 891). 
Malvern Hill, Barbados (PH coll. Febiger 83, 131). 
Mount Hillaby, Barbados (BM coll. Adams GC 3088). 
Newcastle, Barbados (BM coll. Adams F 1236). 

Since we published our previous account of this species (Ross & Sims, 1980), we have been able 
to examine more specimens and to add appreciably to our knowledge of it. The most remarkable 
feature is the inflexed marginal ridge (Fig. 84). This is also present in Dextradonator jeremianus 
R. Ross & P. A. Sims, the other species of the genus, but we know of nothing similar in any other 
genus. 

The stratigraphic range of this species is uncertain. All the specimens that can be firmly dated 
come from the middle Eocene or the middle-upper Eocene boundary, but there can be no 
certainty about the level in the Oceanic Beds from which come the specimens from Bissex Hill, 
Malvern Hill, Mount Hillaby, and Newcastle, and those with no locality detail other than 
'Barbados'. 

In addition to the specimens that we have definitely identified as Dextradonator eximius , there 
is one about which we are uncertain (Figs 82, 83). This comes from the Indian Ocean dredge 
sample that dates from the middle-upper Eocene boundary. It has a diameter of 94 fxm, 
appreciably larger than those about whose identity we are sure, and it differs from them in 
having a marginal ridge that is not turned inwards at its upper edge and that bears two flattened 
spines about 30 ixm tall opposite each other and half-way between the elevations. In all other 
respects it is identical with the specimens that we include in D. eximius with certainty. To decide 
whether it represents a separate species or whether the characteristics peculiar to it are those of 
specimens at the upper end of the size range of D. eximius needs a greater range of specimens 
than those currently available to us. 

Discussion 

As we pointed out in the introduction to this paper, we deal here with a number of genera that do 
not form a closely related group, and hence their affinities and position in the classification 
system of diatoms can only be discussed piecemeal. There are, however, a few general points to 
be made before doing this. As all the genera dealt with in this paper belong to the order 
Biddulphiales, we need to say something about the position and circumscription of this order. 
Very different views on this have been put forward during the last decade. 

This is not the place to consider whether a taxon containing all the diatoms should have the 
level of a division or of a subdivision within the Chrysophyta, but the latter is, in our view, the 
lowest rank to be assigned to it. Some authors, the latest of them Round (1981), have considered 
that, at the next lower level, the diatoms should be divided into three groups: the centric ones, 
the pennate ones without a raphe, and those with a raphe. Others, most recently Simonsen 
(1979), consider that the pennate diatoms with and without a raphe should not be separated at 
this level. This difference of opinion does not, however, affect the circumscription of a taxon 
containing the centric diatoms, which include the Biddulphiales. Hendey (1964), on the other 
hand, rejects the separation of the centric and pennate diatoms, and divides the diatoms into a 
number of groups with the rank of suborder. 

We accept that the centric and pennate diatoms should be separated as distinct taxa at the 
level of class. Whether the pennates should constitute a single class or two, one without and one 
with a raphe, is not relevant here. Within the class Centrophycidae we include an order 
Biddulphiales. We thus agree with Glezer (1979) and Round (1981) about the rank of a taxon 
that includes Biddulphia but excludes Thalassiosira, Coscino discus, Aulacodiscus, and Rhizo- 
solenia, rather than with Simonsen (1979), for whom it is a suborder within the order Centrales. 



290 R. ROSS AND P. A. SIMS 

For both Glezer and Round, the Biddulphiales have a much narrower circumscription than 
that attributed to his suborder by Simonsen; they both include in the order only part of the 
Biddulphiaceae as delimited by Simonsen, to which Glezer adds a few genera included by 
Simonsen in the Eupodiscaceae and the Diatomaceae. We would argue for a much wider 
circumscription of the order than either Glezer's or Round's, but a somewhat narrower one than 
Simonsen's in that we consider that the Lithodesmiaceae should not be included. Also, we are 
not completely certain that either the Chaetoceraceae or the Stictodiscoideae belong in the 
order. There has, in addition, been one important change in the classification of the genera that 
we would definitely include within the Biddulphiales: the establishment of the family Cymatosir- 
aceae Hasle, von Stosch & Syvertsen (1983). We point out below the difficulties in classifying 
into families the remaining genera that we include in the order. 

We regard all the diatoms that we definitely include in the Biddulphiales as having polar 
valves, from unipolar, as in Isthmia Agardh, to multipolar, as in those species of Triceratium 
Ehrenberg for which De Toni (1894) erected the genus Nothoceratium. We consider that the 
ocelli of such genera as Euopodiscus J. W. Bailey and Rattrotyella De Toni indicate that the basic 
morphology of these genera is polar in spite of their circular outline. Also, almost invariably 
throughout the order as we delimit it the internal opening of the labiate process is straight and 
projects as no more than a papilla into the interior of the frustule; it is not borne on a stalk as in 
the Thalassiosirales and Coscinodiscales. The only exceptions of which we are aware are Isthmia 
Agardh (see Navarro, 1981) and Biddulphiopsis von Stosch & Simonsen (1984), in both of which 
the labiate process is shortly stalked internally. Labiate processes of a similar form to that found 
in the Biddulphiales occur in some Melosiraceae, and all those found in pennate diatoms are of 
this type. Another character that is widespread among the Biddulphiales as we delimit them is 
the presence of interstrial pores. The genera in which these occur include not only Biddulphia 
(see Ross & Sims, 1971: pi. 1 fig. 2) and Biddulphiopsis (see von Stosch & Simonsen, 1984), but 
also Hemiaulus (see Ross, Sims & Hasle, 1977: pi. 5 fig. 32, pi. 8 fig. 52), Trigonium (see Ross & 
Sims, 1971: pi. 2 fig. 6), Amphitetras, and Pseudauliscus (unpublished observations). They do 
not seem to occur, however, in the Cymatosiraceae or the Chaetoceraceae, nor outside the 
Biddulphiales. 

Glezer (1979) restricts the order Biddulphiales to genera with poroid areolae, maintaining 
that the type of areola is of primary importance in the classification of the diatoms and that such 
structures as the ocellus and the pseudocellus are polyphyletic (see also Glezer, 1983). She 
accordingly maintains Zygoceros Ehrenberg, a genus with ocelli and loculate areolae (see Ross 
& Sims, 1971), as separate from Odontella Agardh, with ocelh but poroid areolae, whereas 
other authors, e.g. Simonsen (1974), have included the species with loculate areolae in 
Odontella. She places Zygoceros, along with Trigonium Cleve, which has loculate areolae but 
pseudocelli at the apices, in the Coscinodiscaceae, whilst assigning Odontella and Amphitetras 
Ehrenberg, another genus with ocelh and poroid areolae, to the Biddulphiaceae, along with 
Biddulphia Gray which also has poroid areolae but has pseudocelli at the apices. The number of 
characters shared by the genera we include in the Biddulphiales and not present in Coscinodis- 
cus Ehrenberg and its obvious allies indicates that this separation on a priori grounds is unsound. 
All the evidence seems to point to loculate areolae being polyphyletic, and the fact that, when 
areolae become close-packed, they must be of this type for mechanical reasons, reinforces this 
view. More recently, Glezer (1986) has proposed a new monotypic family for Triceratium 
Ehrenberg sensu stricto, and assigned it to the Pyxidiculales. She bases her arguments for this on 
the view that the pseudoloculi of Triceratium are areolae. However, as we showed earlier (Ross 
& Sims, 1971), and Miller & Collier (1978) confirmed, the pseudoloculi of Triceratium are 
formed by a network of external costae laterally expanded at their free edges and the areolae are 
small and poroid. The genus is closely related to Amphitetras, which Glezer (1979) includes in 
the Biddulphiales. 

As well as differing from Glezer on the circumscription of the Biddulphiales, we also do not 
agree with her classification of the genera that she includes within the family Biddulphiaceae. 
She separates the Biddulphiaceae into two subfamilies, one containing those genera with bipolar 
valves, the other those genera with tripolar and multipolar valves, again on a priori grounds. 



GENERA OF THE BIDDULPHIACEAE 291 

There is, however, overwhelming evidence that the change from bipolar valves to tripolar ones 
has happened many times. The range of morphological variation in diatoms of the two shapes is 
very similar and there are many cases where the only differences between species other than the 
number of poles are in the detail of the arrangement and spacing of the areolae, characters that 
in any other context would be regarded as indicating separation at no more than the specific 
level. There are even a number of cases where bipolar and tripolar forms have been treated as 
varieties of the same species, Odontella retiformis (A. Mann) von Stosch (1985) being the most 
recent. Pseudorutilaria adds another example where bipolar and tripolar diatoms must be 
considered as belonging to the same taxon at the generic level. 

There is a nomenclatural point that must also be considered before discussing the rela- 
tionships of the genera described above. The family name Eupodiscaceae has long been used in 
the sense of a family based on the genus Eupodiscus i . W. Bailey, nom. cons., published in 1851. 
However, the family name was published by Kiitzing in 1849 and is based on Eupodiscus 
Ehrenberg, nom. rej., the type of which is Eupodiscus germanicus (Ehrenberg) Ehrenberg, a 
synonym oi Aulaco discus argus (Ehrenberg) A. Schmidt. Eupodiscaceae is thus not the correct 
name for a family within the Biddulphiales. It is also an illegitimate name because Eupodiscus 
Ehrenberg is not only a rejected name but also an illegitimate superfluous substitute for 
Tripodiscus Ehrenberg. 

Solium and Pseudorutilaria resemble each other in most respects. The only basis for generic 
distinction between them is the presence in Pseudorutilaria of the troughs on the central portion, 
the raised edges of which grasp the occluded and labiate processes of the sibhng valve. These we 
regard as a derived feature, and hence we postulate that Pseudorutilaria evolved from an 
ancestor which would, if it were known, be placed in, or very close to, the genus Solium. 

Whilst the fact that the subocelli of Solium and Pseudorutilaria consist of areolae and not 
porelli suggests that their affinities are with Biddulphia and Briggera rather than with 
Amphitetras and Odontella, many of their characters point in the other direction. Their mantles 
are concave, at least in part, as in Odontella and Amphitetras, not vertical as in Briggera and 
most species of Biddulphia, including its type. Anastomosing low external costae, such as occur 
on the elevations and mantles of Solium and Pseudorutilaria, are present throughout on 
Amphitetras but do not occur on Biddulphia or any of its allies. On the other hand, no species 
with a true ocellus has linking spines or internal costae. What is clear, however, is that Solium 
and Pseudorutilaria, in spite of their possession of linking spines, are not to be included in the 
Hemiauloideae sensu Sims (1986). 

The only genus that might be close to Solium and Pseudorutilaria is Monile. It closely 
resembles Pseudorutilaria in its shape of valve, in the many sulci crossing its projections, in the 
concavity of the valve mantle where the valve margin is convex, in the anastomosing costae on its 
elevations, in the arrangement of its poroid areolae, and in the presence of a subocellus. On the 
other hand, its areolae are occluded by cribra, not volae. It is only presence of a subocellus 
rather than a pseudocellus on its elevations , the flattened summits of these , and the ring of radial 
ridges on them that distinguishes it from Biddulphia, although the concavity of the mantle is 
unusual in that genus. The radial ridges on the summits of the elevations have the same pattern 
of arrangement as the linking spines of Strelnikovia R. Ross & P. A. Sims and Keratophora 
Pantocsek, genera which also have well developed pseudocelli. Which of these resemblances are 
indications of affinity and which are parallelisms cannot be decided on the basis of our present 
knowledge. 

The subocellus of Maluina is much smaller than that of Solium, Pseudorutilaria, and Monile 
and its areolae or porelU are much finer and more tightly and uniformly packed. These 
differences are so considerable as to suggest that Maluina may well not have any close affinity 
with those genera. Its elevations bear a smaU number of interlocking linking spines, but these are 
of a different shape from those in Solium and Pseudorutilaria (compare Fig. 67 with Figs 6 and 
13) and it differs from those two genera in virtually every other respect. It seems to be an isolated 
genus and we can offer no suggestions as to its affinities. 

The similarities between Bonea and Strelnikovia are such as to indicate that they are related. 
The many interlocking linking spines and the large pseudocellus are the characters that point to 



292 R. ROSS AND P. A. SIMS 

this most strongly. In Strelnikovia antiqua (Strel'nikova) R. Ross & P. A. Sims and 5. incUnata 
R. Ross & P. A. Sims there is some sign of a vertical division of the pseudocellus at the apex of 
the valve (see Ross & Sims, 1985: pi. 21 fig. 5, pi. 24 fig. 6) and this is more strongly developed in 
Bonea (Fig. 73). We would therefore add Bonea to the group consisting of Briggera R. Ross & P. 
A. Sims, Didadiopsis De Toni, Strelnikovia R. Ross & P. A. Sims, Keratophora Pantocsek, and 
Thaumatonema Greville. As we have already indicated (Ross & Sims, 1985; Sims, 1986), this 
group is much closer to Biddulphia than it is to Hemiaulus, and it should not be included in the 
Hemiauloideae. There are species of Biddulphia that differ from species of Briggera in nothing 
but the lack of interlocking linking spines, and other species that have all the characters of 
Strelnikovia except interlocking linking spines (see Ross & Sims, 1985: 288-290). There is the 
same similarity and difference between Bonea simulans R. Ross & P. A. Sims and Triceratium 
exornatum Greville. The significance of this for the supra-generic classification of the Biddul- 
phiaceae remains obscure, but it does suggest that the transition between diatoms with 
interlocking linking spines and those without them has occurred more than once. There is, 
though, no real indication of the direction in which it happened. 

One of us (Sims, 1986) has recently grouped together in the subfamily Hemiauloideae of the 
Biddulphiaceae the genera Hemiaulus Heiberg, Sphynctolethus Hanna, Pseudaulacodiscus 
Jouse, Ailuretta Sims, Cerataulina H. Peragallo, and Eucampia Ehrenberg. Trinacria Heiberg 
differs from Hemiaulus sensu stricto only in the absence of sulci or internal costae and in the 
number of poles and of labiate processes, and it also clearly belongs in the Hemiauloideae. 
Whether Dextradonator should also be included is more difficult to decide. It differs from 
Hemiaulus in the shape of the valve and of the linking spines, in the absence of sulci or internal 
costae , and in the pattern of areolation . Its very deep , coarsely areolate , entire val vocopula (Fig . 
81) is another point of difference. This is a character that it shares with Abas R. Ross & P. A. 
Sims (1980), which it also resembles in shape of valve and pattern of areolation. We would for 
the present tentatively include both genera in the Hemiauloideae, with the proviso that they are 
not nearly so closely related to the rest of the subfamily as the other genera included in it are to 
one another. 

Our main conclusion must be, then, that the information presented here makes the supra- 
generic classification of the genera that have been included in the Biddulphiaceae and the 
so-called Eupodiscaceae more difficult rather than less. That this is so is probably to be 
attributed to the fact that our knowledge of the Cretaceous and Paleogene diatom floras is still 
inadequate, and will perhaps always be so. 

Acknowledgements 

Our thanks are due to the late Mr A. L. Brigger and to Prof. R. W. Holmes of the University of California, 
Santa Barbara, for material presented to the British Museum (Natural History), which included many 
specimens studied whilst preparing this paper. We also wish to thank Dr Ruth Patrick and Dr C. W. 
Reimer of the Academy of Natural Sciences of Philadelphia, and Dr R. Simonsen of the Alfred-Wegener- 
Institut fur Polar- und Meeresforschung, Bremerhaven, for the opportunity to examine collections in their 
charge. We have once again relied on information about the provenance and dating of samples collected by 
R/V Vema provided by Mrs Margaret Hanna of the California Academy of Sciences and Dr J. Fenner of 
the Geologisch-Palaontologisches Institut und Museum der Universitat Kiel. Type specimens of new 
species were mounted for us by Mr K. D. Kemp of East Brent, Somerset, and Mr S. J. Russell of our 
Department. We also thank the staffs of the photographic unit and electron microscope unit of the museum 
for their assistance. Mr A. Eddy kindly entered the manuscript on a word processor. 

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Description of plates 

Plates 1-11 are SEM micrographs. 

Plates 12-13 are LM micrographs. 

Plate 1. Solium exsculptum Heiberg. Figs 1-3, 5, 6: forma exsculptum, 'Kamischev', presumed to be 

Kamyshlov, Sverdlovsk oblast, U.S.S.R.; Fig. 4: forma, pentagona Jouse, Northern Urals, Tyumen'sk 

oblast, U.S. S.R.; Figs 1, 3, 4: bar line = 10/i,m;Figs2,5, 6: barhne = 2/Lim. 

Fig. 1: valve exterior tilted and displaying four projections with elevations separated from central domed 

area by deep sulci, well-developed marginal ridge (arrowhead), numerous hollow spines (or occluded 



296 R- ROSS AND p. A. SIMS 

processes), and single labiate process (arrow). Fig. 2: same specimen, detail of distal side of elevation with 
subocellus of areolae in radial rows positioned on upper part of elevation, linking spines, and concave 
mantle. Note also costae on mantle and elevation. Fig. 3: oblique view of frustule linked to single valve; 
fragments of girdle bands present. Fig. 4: oblique view of valve exterior of 5-polar form with concave 
mantle, sulci separating projections from central portion which has irregularly scattered areolae and an 
off-centre single labiate process. Fig. 5: distal side of linked elevations with clearly-defined subocelli 
apposed and single rows of poroid areolae between anastomosing costae. Note each areola is bordered by a 
rim with a single small spine and occluded by a vola. Fig. 6: proximal face of tips of two elevations linked by 
expanded and interlocking linking spines. 

Plate 2. Pseudorutilaria monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni, Division 
Hill, Otago, New Zealand. Figs 7-9, 11: bar line = 20 )u,m; Figs 10, 12, 13, 14: bar hne = 3 /xm. 
Fig. 7: girdle view of two valves linked at elevations. Note central circular domed area, also regular rows of 
spines arising from marginal ridge between central portion and each elevation. Figs 8,9: valve exterior with 
almost circular central portion and projections with crenulate margins gently tapering from central portion 
to each pole. Fig. 10: central portion with entire margin, sparse and scattered poroid areolae, irregular ring 
of troughs with raised margins (one broken occluded process (arrow) clasped within trough) mostly 
alternating with the fractured occluded processes (arrowhead). Fig. 11: two linked valves, oblique view of 
valve interior showing transapical costae positioned at each indentation of the valve margin. Fig. 12: valve 
interior with opening from single labiate process. Fig. 13: interlocking linking spines at tips of linked 
elevations on proximal sides. Fig. 14: oblique view of two linked elevations with a subocellus positioned 
just beneath each tip, linking spines on the proximal faces and solid spines arising from marginal ridge. 
Plate 3. Pseudorutilaria clavata R. Ross & P. A. Sims. Figs 15, 16, 18-23: south-western Atlantic, 
Falkland Plateau, 51° OS'S, 54° 22' W, 1525 m depth, Vema cruise 17, core 107, 50 cm. Fig. 17: 
south-western Atlantic, Falkland Plateau, 53° 01' S, 52° 52' W, 2880 m depth, Vema cruise 18, core 104, 
330 cm. Figs 15-17, 19, 20: bar hne = 20 /im; Figs 18, 21-23: bar line = 5 /xm. 

Figs 15-17: exterior views showing valve outline with almost circular central portion, gibbous portion on 
either side with each projection tapering to clavate rounded apices. Note marginal spines on marginal ridge 
and undulate valve face. Fig. 18: girdle view, two linked valves at centre, each with occluded processes and 
single labiate process (arrow) clasped within troughs with raised margins (two arrows) of opposing valve. 
Note girdle band with finely poroid areolae. Figs 19, 20: valve interior with four transverse sulci visible, 
positioned between central portion, gibbous portions, and projections. Note valve margin is entire, also 
note opening from single labiate process on fig. 20. Fig. 21: central portion with ring of occluded processes 
(broken at base), base of labiate process (arrow), also troughs with raised margins, one enclosing process 
of sibling valve. Fig. 22: distal side of low elevation with subocellus and four linking spines, one broken at 
base. Fig. 23: girdle view showing interlocking linking spines at tips of proximal faces of low elevations, 
apposed subocelli clearly defined, also flattened spines arising from marginal ridge. Note also deep furrow 
between mantle and hyaline marginal band, also anastomosing costae. 
Plate 4. Pseudorutilaria nodosa R. Ross & P. A. Sims. 

Figs 24-27, 29, 32: 'Lamont E8'; Figs 28, 33: south-western Atlantic, Falkland Plateau, 53° 01' S, 52° 52' 
W, 2880 m depth, Vema cruise 18, core 104, 330 cm. Figs 24-28, bar line = 20 /^m; Figs 29-33, bar Hne = 5 
/u,m. 

Fig. 24: oblique view of two linked valves showing valve interior with numerous transverse costae. Fig. 25: 
oblique view of valve exterior showing undulate valve face, domed central portion, distinct marginal ridge 
and vertical mantle with deep furrow above narrow marginal band. Figs 26, 27: same specimens lying flat, 
displaying distinctive valve outline. Fig. 28: specimen with straight, not curved valves. Note position of 
gibbous portions and arrangement of internal costae identical with those above. Fig. 29: central portion, 
exterior view, with ring of fractured occluded processes alternating with troughs with fractured margins, 
also scattered poroid areolae. Fig. 30: Hnked elevations with subocelli on distal side and linking spines on 
proximal face, also long solid spines arising from marginal ridge. Fig. 31: single elevation with eroded 
linking spines. Fig. 32: two linked valves, central portion, with external openings from two labiate 
processes visible (arrows), one from each valve inclined and lying on opposing valve margin. Note also 
raised margins of troughs and indentation at base of mantle . Fig . 33 : interior view , central portion , showing 
opening from single labiate process and transapical costae. 

Plate 5. Pseudorutilaria hannai R. Ross & P. A. Sims. South-western Atlantic, Falkland Plateau, 53° 01 ' S, 
52° 52' W, 2880 m depth; Vema cruise 18, core 104, 330 cm. All figures are of the same specimen. Figs 34, 
35: bar line = 30 )u.m; Figs 36-38: bar line = 5 /am. 

Fig. 34: two linked valves, interior view with central circular portion defined by transverse costae and 
curved projections regularly crossed by costae. Fig. 35: specimen in almost girdle view showing undulate 
margin and spines from marginal ridge of both valves alternating along complete length of valve. Fig. 36: 



GENERA OF THE BIDDULPHIACEAE 297 

view at pole with interlocking linking spines on proximal face of each low elevation, subocellus missing on 
distal side and indentation or deep furrow between concave mantle and marginal hyaline band. Fig. 37: 
exterior opening of labiate process from upper valve (arrow) positioned between marginal spines. Fig. 38: 
central portion with slit-like opening from single labiate process (arrow). Note also scattered areolae and 
furrow between mantle and valve margin in central portion. 
Plate 6. Pseudorutilaria incompleta R. Ross & P. A. Sims. 

Figs 39-46: south-western Atlantic, Falkland Plateau, 51° 08' S, 54° 22' W, 1525 m depth, Vema cruise 17, 
core 107. Fig. 47: south-western Atlantic, Falkland Plateau, 47° 45-7' S, 57° 38-5' W, 3650 m depth, Conrad 
cruise 12, core 237. Figs 39-41: bar line = 20 /im; Figs 42-44: bar line = 2 )u,m; Figs 45-47: barUne = 5/xm. 
Figs 39, 40: valve exterior of separation valve and valve within chain displaying distinctive valve outline 
with crenulate margin, sulci producing undulate valve surface and the pentagonal central portion. Fig. 41: 
two linked valves tilted to show valve interior with sulci positioned between each crenulation and a large 
pentagonal central portion. Fig. 42: separation valve at pole with subocellus positioned just beneath tip of 
low elevation and single large spine on proximal margin. Fig. 43: distal side of linked elevations with 
discrete subocellus beneath tip of each elevation. Fig. 44: girdle view of same specimen with discrete 
subocelli and interlocking linking spines, areolae with vela, also marginal spines interdigitating. Fig. 45: 
central portion of separation valve with single straight labiate process, hyaline circular area surrounded by 
low flange-like costa, also scattered poroid areolae. Fig. 46: central portion, valve exterior with base of 
single labiate process (arrow), occluded processes and eroded troughs with raised margins arranged in 
circle. Fig. 47: central portion, valve interior, with opening from single labiate process, crenulate margin 
and internal sulci. 

Plate 7. Pseudorutilaria hendeyi R. Ross & P. A. Sims. 

Figs 48, 49, 53: south-western Atlantic, Falkland Plateau, 53° Ol'S, 52° 52' W, 2880 m depth, Vema cruise 
18, core 104, 330 cm. Figs 50-52, 54: south-western Atlantic, Falkland Plateau, 51° 08' S, 54° 22' W, 1525 m 
depth, Vema cruise 17, core 107, 50 cm. Figs 48, 50: bar line = 20 /u,m; Figs 49, 51-54: bar line = 5 /am. 
Fig. 48: oblique view of triangular valve with central area defined by three sulci, long projections with 
undulate surface and rounded apices. Fig. 49: central portion fractured at centre but surrounded by 
remains of occluded processes, labiate processes and troughs. Fig. 50: interior view, two linked valves with 
transverse sulci proximally and costae distally across each projection and crenulate valve margin. Fig. 51: 
interior view, central portion, with openings from two labiate processes, scattered poroid areolae and 
patch of porelli. Fig. 52: fractured specimen, two linked valves, view looking into valve centre with 
occluded processes (arrow) and troughs with raised margins (arrowheads). Note marginal spines interdigi- 
tating. Fig. 53: single projection with low elevation. Exterior view showing position of linking spines on 
proximal face. Fig. 54: two linked valves, polar view of projection with low elevations, remains of 
well-defined subocelli and internal costae on upper valve. 

Plate 8. Monile laurentii R. Ross & P. A. Sims. St. Laurent-la- Vernede, Gard, France. Figs 55-60: bar line 
= 25 /Am; Figs 61-64: bar line = 5 ixra. 

Figs 55, 56: single valves, exterior view, showing sinuous valve outline, domed central portion, sub-clavate 
apices and elevations with flattened summits. Fig. 57: oblique view, valve exterior, with vertical mantle on 
which there are three central and a polar indentation. Figs 58-59: interior view, single valves with 
transapical sulci positioned at each indentation of valve margin. Fig. 60: frustule with attached girdle band. 
Upper valve with undulate valve face, openings from four labiate processes on central portion and 
flattened summits to short, squat elevations. Fig. 61: central portion with exterior openings from four 
labiate processes projecting as fractured hollow spines, poroid areolae each with slightly raised rim with 
one or two small solid spines and occluded by cribrum. Fig. 62: interior view, central portion, with openings 
from four labiate processes. Fig. 63: proximal face of elevation with flattened summit encircled by marginal 
row of low teeth, areolae with cribra and interstrial pores. Fig. 64: distal side of elevation, expanded above, 
flattened summit bearing ring of low radial teeth on margin and subocellus on summit and expanded tip. 
Note anastomosing costae on elevation. 

Plate 9. Maluina centralitenuis (R. Ross & P. A. Sims) R. Ross & P. A. Sims. Figs 65-70: south-western 
Atlantic, Falkland Plateau, 51° 08' S, 54° 22' W, 1525 m depth, Vema cruise 17, core 107, 50 cm. Fig. 65: 
bar line = 20 /tm; Figs 66-70; bar line = 5 /am. 

Fig. 65: two linked valves with cylindrical elevations, vertical mantles with strongly developed pseudo- 
antiligulae at apices and domed centres. Fig. 66: detail of mantle with single row of poroid areolae and 
pseudo-antiligula, central domed area with radial rows of poroid areolae and single labiate process. Fig. 67: 
proximal face of cylindrical elevations with linking spines, expanded above and interlocking. Fig. 68: distal 
side of elevations, each with small subocellus at its tip. Note scattered areolae. Fig. 69: valve interior with 
large, slit-like opening from single labiate process, rows of fine areolae radiating from valve centre to 
margin of dome, also on outer valve face but sparse on vertical mantle. Fig. 70: valve exterior with small 



298 R. ROSS AND P. A. SIMS 

circular hyaline central area, rows of fine poroid areolae radiating to base of domed centre, also opening 
from single labiate process projecting as a small tubular spine. 

Plate 10. Bonea simulansK. Ross & P. A. Sims. 'Carlovo',U.S.S.R. Figs71,72,74: bar line = 20 /xm; Figs 
73, 75-77: bar line = 3 /xm. 

Fig. 71: valve exterior, triangular valve with low elevations with flattened summits, marginal ridge between 
elevations bearing large spines, also vertical mantle. Fig. 72: valve interior showing weakly folded valve 
face with opening from single central labiate process. Fig. 73: elevation with summit surrounded by 
complete ring of expanded solid spines, those on the distal margin fractured. Note pseudocellus on summit 
and tip of elevation. Fig. 74: girdle view of frustule linked to two valves with two girdle bands present. Note 
pseudocellus on distal side of each elevation, also the interdigitating marginal spines. Fig. 75: distal side of 
elevations linked by numerous small expanded spines which interlock with those of the apposing elevation. 
Fig. 76: poroid areolae occluded by rotae with c. nine spokes, also single interstrial pore. Fig. 77: occlusion 
to smaller poroid areolae, also two interstrial pores. 

Plate 11. Dextradonator eximius (Grunow) R. Ross & P. A. Sims. Indian Ocean, 10° 25' S, 63° 15' E, 3115 
m depth, dredge sample DODO-123-D1. Figs 78-83: bar line = 20 /nm; Figs 84-85: bar Hne = 10 /u,m. 
Identity of specimen shown in Figs 82-3 doubtful (see p. 289). 

Fig. 78: two valves Unked, upper with hyahne band (pleura) attached. Note linking mechanism involving 
two flattened and acute Unking spines partially encircling elevation of sibling valve. Fig. 79: oblique view, 
same specimen, with hyaline marginal ridge turned inwards between elevations (arrows). Fig. 80: linked 
circular valves with broad vertical mantles, upper with attached valvocopula, lower with hyaline mantle 
and marginal ridge turned inwards. Fig. 81: single valve with attached valvocopula, cylindrical elevations 
each with two fractured linking spines. Fig. 82: Hnked valves, girdle view. Note two spines from each valve 
projecting beyond the girdle bands. Fig. 83: oblique view, valve face with small hyaUne central area, 
radiating rows of fine poroid areolae extending to marginal ridge and shghtly beyond base of elevations, 
also two long spines arising from marginal ridge. Fig. 84: distal side of elevation with few scattered areolae 
at tip, also circular valve face with marginal ridge turned inwards. Fig. 85: valve interior, showing central 
hyahne area, rows of fine poroid areolae radiating from centre, also broad valve margin. 
Plate 12. Figs 86-88: Solium exsculptum Heiberg. 'Kamischev' presumed to be Kamyshlov, Sverdlovsk 
oblast,U.S.S.R. 

Fig. 86: fo. exsculptum BM 81304: frustule with valves linked on either side, girdle view, 86-5 x 32 /u,m. Fig. 
87: fo. pentagona Jouse, BM 65833: 36 /xm diameter. Fig. 88: fo. exsculptum BM coll. Adams TS748: full 
length one side, 67 /am. Fig. 89: Pseudorutilaria monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt 
ex De Toni, BM 63397: Oamaru, New Zealand: length of specimens a: 136 ju-m, b: 97 /itm, c: 130 /am. Figs 
90, 91: Pseudorutilaria clavata R. Ross & P. A. Sims, BM 81136: south-western Atlantic, 51° 08' S, 54° 22' 
W, 1525 m depth; Vema cruise 17, core 107, 175 cm; Fig. 90: girdle view, two linked valves; apical axis 156 
/am; Fig. 91: Holotype; apical axis 95 /am. Fig. 92: Pseudorutilaria nodosa R. Ross & P. A. Sims, Holotype, 
BM 81138: south-western Atlantic, 51° 08' S, 54° 22' W, 1525 m depth; Vema cruise 17, core 107, 50 cm; 
apical axis 146 /am. Fig. 93: Pseudorutilaria hannai R. Ross & P. A. Sims, Holotype, BM 81300: 
south-western Atlantic, 53° 01' S, 52° 52' W, 2880 m depth; Vema cruise 18, core 104, 330 cm; apical axis 
152 /am. 

Plate 13. Figs 94-95: Pseudorutilaria incompleta R. Ross &. P. A. Sims, BM 81139: south-western 
Atlantic, 51° 08' S, 54° 22' W, 1525 m depth; Vema cruise 17, core 107. Fig. 94: Holotype; apical axis 98 
/am. Fig. 95: two linked valves; apical axis 80 /am. Fig. 96: Pseudorutilaria hendeyi R. Ross «fe P. A. Sims, 
Holotype; BM 81142: south-western Atlantic, 51° 08' S, 54° 22' W, 1525 m depth; Vema cruise 17, core 
107, 170 cm; apex to apex = 180 /am. Figs 97-98: Monile laurentii R. Ross & P. A. Sims, St 
Laurent-la- Vernede, Gard, France. Fig. 97: Holotype, BM coll. Adams H941, specim. I, 12: apical axis 
220 /am. Fig. 98: BM 81303: apical axis 228 /am. Figs 99-100: Bonea simulans R. Ross & P. A. Sims. 
'Carlovo', U.S.S.R.;Fig. 99: Holotype, BM 81301: length of side 119 /am. Fig. 100: BM 81302: girdle view; 
length of side 140 /am. 



PLATE 1 



299 




300 



PLATE 2 




PLATE 3 



301 




302 



PLATE 4 




PLATE 5 



303 




304 



PLATE 6 




PLATE 7 



305 




306 



PLATE 8 




y^ 



PLATE 9 



307 




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PLATE 10 




PLATE 1 1 



309 




310 



PLATE 12 




PLATE 13 



311 




\ 



British Museum (Natural History) 

An Enumeration of the Flowering Plants of 
Nepal 

H. Hara, W. T. Stearn, A. O. Chater & 
L. H. J. Williams 

Vol. 1. The Gymnosperms and Monocotyledons 

1978, 275 X 215mm, 154pp, 7 figs. Paperback. 
565 00777 7 £30.00 

Vol. 2. The Dicotyledons 

(Part). 1979,220pp. 
565 00810 2 £30.00 

Vol. 3. The Dicotyledons 

(Part). 1982, 226pp,l fig. 
565 00854 4 £35.00 



Titles to be published in Volume 16 



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

Ttie lichen genus Ramalina in Australia 

By G. N. Stevens 

An annotated hst of vascular plants collected in the valleys south of 
Mt Everest 

By G. Miehe 

Further genera of the Biddulphiaceae (diatoms) with interlocking linking spines 

ByR. Ross&P. A. Sims 



Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk 
Printed in Great Britain by Henry Ling Ltd, Dorchester