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Bulletin of the patorres
GENER
British Museum (Natural History)
Botany series’ Vol 16 1987
British Museum (Natural History)
London 1987
Dates of publication of the parts
Nol. : ; : : ‘ ; . ‘ : ; 29 January 1987
No2s:, : F : ; ; : ‘ : : : 25 June 1987
No3. : é : : : ‘ : : ‘ : 27 August 1987
No4 . : ; : : : : ; : : 26 November 1987
ISSN 0068-2292
Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset
No 1
No2
No 3
No 4
Contents
Botany Volume 16
Studies in the genus Hypericum L. (Guttiferae)
7. Section 29. Brathys (part 1)
N. K. B. Robson ; :
The lichen genus Ramalina in Australia
G.N. Stevens .
An annotated list of vascular plants collected in the valleys south of
Mt Everest
G. Miche.
Further genera of the Biddulphiaceae (diatoms) with interlocking
linking spines |
R. Ross and P. A. Sims
Page
225
269
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Bulletin of the
British Museum (Natural Bio)
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
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Bot.)
© Trustees of the British Museum (Natural History), 1987
The Botany series is edited in the Museum’s Department of Botany
Keeper of Botany: Mr J. F. M. Cannon
Editor of Bulletin: Mr J. R. Laundon
Assistant Editor: Dr A. J. Harrington
Editor’s Assistant: Miss M. J. Short
<< SH MUSED
v YBLICA MONS
ISSVED
Lig
if XN
ISBN 0 565 08014 8 Hs =
ISSN 0068-2292 ies ams ees a * Ts Botany series
* Pi Se % Vol16No 1 pp 1-106
British Museum (Natural Hist py WALES OFF a Sy Ss he
Cromwell Road oe TURAL His SF0x
London SW7 5BD Issued 29 January 1987
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
A AEs Desk | DET OANA TOS PRR OC ARPT Ca OAS THERON PETS PET ONT EAE Ee REIET OST ey Ae 1
RS ICRC GI CHOIG rosa sc ran ch Saco eke tene eddy Veoh eve edad e coney ta saueseboe ye ceevans diol Snassesuaeestorrs 2
WIDEB OLOGY 5. cries seer tat esse neers dousso- ca iea carwacsetler shun coene aw on opesseanak Hine eed des the 3
ERP UASCAVES: Mireles h tanks ce scrut oo tata nese ih doane coer or Poe aecdee ane tae ters tetas 5
RU PMEOT SCONCES aa tre ecred seen nccan Usnechtumde noany sas eine cdua sk Vin soe eee b
Wey PIG WETS ain SUNS oso ny usa noe sen nee hohe Ca sdreane aint de ating a ica ne itconst 5
ESVCOIOG Ms IY DIIIS o25 esc lays ots suey roo etay cae oes cattechac scree asansetsaitar aie? ol
PPISUTIOULIOM AIO VOM, o, 00 sate rss onea rts Sa one onesies nase va cseaweeeadedeadiy bees} it
CO) WV IGE CIS IUNICHIONS Cort. onersescccrnts oes nce) car is iis dost cats Soe oa ee tae oad coerce ei
(b) Species with leaves persistent or deciduous near the base
SDR IS 25 ee aay eee Rie eds IYER A A teed 9
(c) Species with leaves with sheathing leaf-bases or marginal
Secrenons(Spps 24226. Ola oid het a vetet eck cmt eer smuaien: 10
(d) Species with incurved-cucullate leaves but no marginal
SECTELONSI( SPP 2h SU) ecu cst siies oo sate ee ales ce ee cae ma wma eaannd dee 10
(e) Species with sessile leaves deciduous above the base but
notincurved-cucullate (Sop; 31.0: —48 )s.25;.< foc-anwns cos cc tetas sees -kaceen 11
(f) Species with pseudopetiolate leaves deciduous above the base
but not incurved-cucullate (Spp. 31p.p., 49-60) «20.0.0... eee ce ec ee eee ee es 12
em Phat LOLOL on URES meee oie Pere Sea ayer e Pon ie in Pee tee aM REET Cate ESSN 12
eRe MNO WICCEEMIONES serit cau tnsy aac esier etn ere ceed As sous cane Teneo au a euta mes toes ea teaaoree 103
EN ong Lo} £1 2h coed SCRE Eee Oe re RNS tae eon AOE PEE ACRE PL RET OEN ES RO ee ee 104
Pic AnsVS CCA UIC MINOR oo ca. ors cassie tere cso see na Ren Ce A artist Rumen ad | ps arean aN ae eam auttae 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 Santandér/César), 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. maguirei N. Robson (Ecuador: Azuay, Loja, El Oro), H. gleasonii N. Robson (Colombia:
Norte de Santandér), H. prietoi N. Robson (Ecuador: Azuay), H. cassiopiforme N. Robson (Peru:
Amazonas), H. magdalenicum N. Robson (Colombia: Magdalena; Venezuela: Zulia, Mérida), 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 Santandér/
Bull. Br. Mus. nat. Hist. (Bot.) 16 (1): 1-106 Issued 29 January 1987
2 NORMAN K. B. ROBSON
César), H. marahuacanum N. Robson (three subspecies): subsp. marahuacanum (Venezuela: Amazo-
nas), subsp. strictissimum N. Robson (Colombia: Boyaca; Venezuela: Mérida), subsp. chimantaicum N.
Robson (Venezuela: Bolivar), H. horizontale N. Robson (Colombia: Norte de Santandér, Santandér), H.
jaramilloi N. Robson (Costa Rica; Colombia: Magdalena, César/Norte de Santandér), 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 1—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 H. piriai Arechav. (including H. hilaireanum L.B.Sm. = H. tenuifolium St. Hil. non
Pursh) represents a development from Sect. Brathys in which the flower has apparently become
adapted for specialized insect pollination in a manner partially analogous to those of sects Elodes
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 (H. japonicum Thunb. ex Murray) and Africa and
Madagascar (H. /alandii group). Until now this section, which includes both shrubs and herbs,
has not been recognized as distinct and therefore has no name. It does, however, include the
type species of the genera Knifa Adanson (H. mutilum L.) and Tridia Korthals (H. japonicum
Thunb. ex Murray), both of which were treated as series of Sarothra sect. Spachium by Kimura
(1951). I have therefore chosen Knifa as the epithet for this new section:
Hypericum section Knifa (Adanson) N. Robson, stat. nov.
Knifa Adanson, Fam. Pl. 2: 44, 541 (1763).
Tridia Korth. in Tijdschr. Natirl. Gesch. Physiol. 3: 17(1836).
Sarothra sect. Spachium series Knifa (Adanson) Y. Kimura in Nakai & Honda, Nova FI.
Japonica 10: 233 (1951) ‘Kniffa’.
Sarothra sect. Spachium series Japonica Y. Kimura, tom. cit.: 233 (1951).
Type: H. mutilum L. (Y. Kimura, 1951, lectotype).
The other section, comprising shrubs, suffrutices, and herbs, is centred in south-eastern Brazil
and extends in area to Uruguay, northern Argentina, and Peru and thence northward along the
* For part 1 see Robson (1977); for part 2 see Robson (1981); for part 3 see Robson (1985).
THE GENUS HYPERICUM L. 3
Cordillera to Venezuela. It also occurs in Australasia and SE Asia (H. gramineum Forster f.).
The type of Kimura’s Sarothra sect. Trigynobrathys (H. myrianthum Cham. & Schlecht.)
belongs to this section (although that epithet is not especially appropriate to the concept adopted
here), as well as the type of sect. Brathys subsect. Connatum R. Keller (H. connatum Lam.):
Hypericum section Trigynobrathys (Y. Kimura) N. Robson, comb. nov.
Sarothra section Trigynobrathys Y. Kimura in Nakai & Honda, Nova Fl. Japonica 10: 233
(1951).
Type: H. myrianthum Cham. & Schlecht. (Sarothra myriantha (Cham. & Schlecht.) Y.
Kimura).
The characters by which species in these three sections may be recognized can be summarized
as follows:
1. Leaf lamina with short, sometimes thick basal insertion, pairs free or united by interfoliar ridge
or very rarely more broadly united; inflorescence-branching usually pseudo-dichotomous or
MONOCHAS A door oe oe, ote Pace cule iatened te eevee ye edtraa st wo tot eee 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\, 24.0. 2..4/:..39: fuk: seteeccesacaceuvanaececesuewsesee 2
2. Leaves narrowed at the base or wholly narrow, united by interfoliar ridge, with basal vein
single, venation pinnate or 1-nerved, or, if (secondarily) amplexicaul and 3—5-nerved (H.
majus and H. lalandii groups p.p.), then midrib branches absent of weak.... sect. 30. Knifa
Leaves amplexicaul with 3—7 basal veins and midrib clearly branched, the leaf pairs usually
GOCUITENLOK SODIGHIMICS perloliate <2. .ss0425.so0eascag entanssucsion ters sect. 31. Trigynobrathys
Morphology (Fig. 2)
The species in sect. 29 Brathys vary from small trees (up to 4 m in height) to prostrate shrublets
or wiry herbs. They all lack dark glands completely and have no resin glands on the lower surface
of the leaf (the latter are common in sect. 3 Ascyreia but not in sect. 1 Campylosporus), and there
are no fringing glands on leaves, sepals, and petals. It appears that all have marginal punctiform
leaf glands (although these are often very small), and the laminar glands in the leaves are also
always punctiform. In the sepals and petals, however, the laminar glands vary from wholly linear
to wholly punctiform.
(a) Leaves
The reduction of the laminar leaf glands from lines to dots is not here associated with the
development of reticulate venation, as it isin sect. 1 Campylosporus (Robson, 1981: 80, Fig. 11).
In the species of sect. Brathys with the most primitive characters (1 H. terrae-firmae) the
venation is parallel and open, even though the laminar glands are punctiform. The primary
venation remains open in nearly all the woody species, thus allowing reduction to 1-nerved
leaves to occur in several parts of the section. Only in some of the relatively primitive species,
e.g. 24 H. magniflorum, is there a clear tertiary reticulum.
In sect. Brathys the ultimate fate of the leaves has proved to be of major taxonomic
importance. In the most primitive state (found in 1 H. terrae-firmae and 2 H. styphelioides) the
leaves are free and completely deciduous, leaving a rhombic leaf scar. In all other species they
are wholly or partly persistent, the persistent part being eventually shed with the cortex of the
stem internode; and the leaf bases are usually united by an interfoliar ridge. The ultimate fate of
these leaves depends on their thickness and size. Where they are thick and long, they break off
without fading, leaving a perceptible part of the base (25 H. gleasonii to 61 H. cymobrathys
except 54 H. millefolium). Where they are thin and long, they tend to droop and wither on the
stem (3a H. phellos subsp. phellos in part, 10 H. simonsii, 11 H. papillosum, and 24 H.
magnifiorum). Where these thin leaves have (in evolution) become smaller or narrower and
thicker, they either fall at the apparent base (3 H. phellos and 4 H. garciae) or persist and wither
(Spp. 5-9, 12-23), tending to become secondarily appressed if the base is narrow (e.g. 8 H.
stenopetalum) and depressed if it is broad (e.g. 16 H. cuatrecasii). Finally, in 54 H. millefolium
NORMAN K. B. ROBSON
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THE GENUS HYPERICUM L. 2s
the leaves are very small and remain attached to the stem, falling with it either when it
disarticulates at the nodes or with the cortex.
In those taxa in which the leaves fall at the apparent base, the true base has become united
with the stem, surviving only as a protuberance (e.g. in 4 H. garciae). Such leaves can be
distinguished from the wholly deciduous leaves of H. terrae-firmae and H. styphelioides by the
absence of a true leaf scar.
The order of these leaf characters in evolution would appear to have been: wholly deciduous —
wholly persistent — partly deciduous — wholly persistent again (Spp. 5—9 and 62-80), see Fig. 2.
The leaf base mostly remains parallel-sided or only slightly narrowed before widening to form
the interfoliar ridge; but in one group (31 H. pimeleoides in part and Spp. 49-53, 55, 59 in part) it
narrows perceptibly to form a pseudopetiole. Where the lamina has narrowed to the width of the
pseudopetiole, however, members of this group cease to appear petiolate and have to be
described as (secondarily) sessile (Spp. 54, 56-59 in part).
(b) Inflorescences
The inflorescence in woody members of sect. Brathys is nearly always 1-flowered (the primitive
state in the genus). Only in a few of the more primitive species (e.g. 3 H. phellos, 26 H.
mexicanum, and 61 H. cymobrathys) is there any development of cymose branching; and even
then it is not usually constant and never elaborate. Apart from this relatively weak tendency to
produce monochasia, the inflorescence in woody species is always strictly a synflorescence with
variation resulting from different types and aggregations of 1-flowered branches. The basic form
is the pseudo-dichotomy, i.e. the development of equal foliate branches from the axils of the leaf
pair immediately below the flower. This is the only type of acropetal inflorescence development
in the section; but the upper lateral branches are also often floriferous, and the relative
frequency of these two modes of branching can be of taxonomic value. Thus in 10 H. simonsii, for
example, the flowers are solely on lateral branches, whereas in 21 H. martense the majority of
the pulviniform plant consists of a pseudo-dichotomous synflorescence, with the branches of the
latest pseudo-dichotomies bearing lateral flowering branches (Fig. 8).
The development of a suffruticose or herbaceous habit (6 H. piriai and Spp. 62-80) is
accompanied by the elaboration of monochasial cymose inflorescences.
(c) Flowers and fruits
In section Brathys the perianth is normally pentamerous, with the sepals appressed to the petals.
These are longer than the sepals and usually spreading (‘flowers stellate’), but in small flowers
they may remain ascending (‘flowers obconic’). The sepals vary in width and hence in the
number of nerves, which may be of taxonomic significance, as may also be the aspect (plane to
incurved-cucullate). Variation in petal shape and size may help to separate related species; but
the degree of development of the apiculus, though striking, is not discontinuous enough to be of
taxonomic value.
The number and size of the stamens are correlated in general with flower size as a whole, but
the woody species with the fewest stamens (35 H. decandrum, which can have as few as 5) does
not have the smallest flowers. That distinction belongs to 42 H. prostratum, in which they can be
as small as 4 mm in diameter. Among the herbs H. gentianoides can have as few as 5 stamens and
also has the smallest flowers.
The styles vary in number from 5 (H. terrae-firmae, H. styphelioides) to 3, 4 as usual being an
apparently unstable number, not characteristic of any one species. The majority of species have
3 styles, but in three advanced woody species this trend has been reversed so that an increased
number (4—5) of styles is associated with very small flowers (41 H. juniperinum = H. brathys), 42
H. prostratum, and 60 H. selaginella). The form of the styles (outcurving, incurving, etc.) may be
taxonomically useful. The stigma is primitively narrow but has frequently become broader,
usually in a capitate form; in one group (Spp. 51-53) the style apex is clavate. The ovary and
capsule remain enclosed in the perianth except when the flower is open, and so it is not always
easy to observe them in herbarium specimens. Nevertheless, their shape varies and can be
utilized taxonomically.
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THE GENUS HYPERICUM L. yi
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 H. 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 (J. & C. Taylor 11757 (NY)) and looks like H. irazuense Xx
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
Campylosporus, 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, 1.e.
until the Lower Cretaceous (Turonian—100m yrs BP) (Raven & Axelrod, 1974; Owen, 1976;
Howarth, 1981); but whether it had split into two before or after reaching that region is
uncertain. From the evolutionary divergence pattern of Hypericum as a whole (Robson, 1981:
Fig. 73; Robson, 1985; Fig. 1), it seems likely that the first split occurred in Africa.
For a temperate genus such as Hypericum that is confined in tropical latitudes to high
altitudes, the ranges of the Cordillera act as highways of dispersion to which the species are
restricted and along which they migrate and differentiate. It is therefore often possible to trace
presumed evolutionary lines (clades or, more accurately, morphoclines) in sequence along one
or more of the Andean ranges. If the variation is continuous, as it is for examples in 20 H.
laricifolium, no infraspecific taxa can be recognized, even though the forms at the extreme ends
of the range of this species (‘H. /aricoides’ in Venezuela and ‘H. acerosum’ in Peru) look very
different from the primitive form from northern Ecuador.
If the variation becomes discontinuous, on the other hand, one finds a geographical
replacement series of taxa each more advanced (apomorphic) than the one before. In such a
clade it is frequently difficult or impossible to discover any apomorphic character in the less
advanced taxon that would enable one to conclude that it had continued to evolve independently
subsequent to its isolation from the more advanced one. In such clades, therefore, where it is
difficult to believe in the Hennigian maxim that one taxon is automatically altered when it gives
rise to another (i.e. that at each speciation the cladogram must dichotomize), I have depicted the
relationship (Figs 1-3) by a line, not a dichotomy.
(a) Wide disjunctions
The majority of the primitive species in sect. Brathys occur in, or are confined to the mountains
near, the Colombian-Venezuelan border area or to adjacent areas in Boyaca (Colombia) and
Mérida, Tachira or Trujillo (Venezuela); but the most primitive (plesiomorphic) characters are
found in the species from Belize (1 H. terrae-firmae) and Cuba (2 H. styphelioides), species that
are so closely related that they have sometimes been united under the latter name, e.g. by
Standley & Williams (1961). The question therefore arises as to whether (i) the original
members of sect. Brathys migrated and differentiated southward from Belize/Cuba to Col-
ombia/Venezuela before isolation of the two groups, or (ii) the differentiation took place
NORMAN K. B. ROBSON
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THE GENUS HYPERICUM L. 9
northwards from South America to the Greater Antilles before isolation, or (iii) a uniform
population was split and the two groups evolved in isolation (Fig. 3a/b). What is clear is that this
distribution gives strong support to the theory that the first land link between South America and
North American was by way of the Greater Antilles (Rosen, 1976, 1985; Robson, 1981: 213;
Melville, 1981: 247).
The remaining evolution of the section has been wholly within South America, except for: (i)
four species that have reached Costa Rica/Panama after the establishment of the second land
link (Isthmus of Panama), (7 H. irazuense, 38 H. costaricense, 49 H. jaramilloi, and 50 H.
cardonae), (ii) three shrubby species in Hispaniola (52 H. ekmanii, 53 H. pycnophyllum, and 54
H. millefolium), and (iii) the derivatives of 61 H. cymobrathys, which occur throughout Central
America and the Greater Antilles and extend their area of distribution into eastern U.S.A. The
three Hispaniolan shrubs terminate a clade that originates with two of the Costa Rican species,
which also occur in the basic area of the Colombian- Venezuelan border (49 H. jaramilloi, 50 H.
cardonae) (Fig. 3, clade xxviii). Occupying an intermediate position is 51 H. caracasanum of
northern Venezuela, of which the more advanced subspecies (51b subsp. turmiquirense) is found
at the north-eastern extreme of the Andean range and is also morphologically close to 52 H.
ekmanii. These data suggest that the ancestor of the Hispaniolan species reached that island
either directly by long-distance dispersal or via the Lesser Antilles, which, according to the
above mentioned theory, would by then have arisen as a result of volcanic action; but overland
access before the separation of the Greater Antilles from South America cannot be ruled out. A
three-fold origin of Hispaniola from parts previously associated respectively with Yucatan,
eastern Cuba, and the Bahamas, such as has been proposed by Rosen (1985), would make the
overland route less likely (see also p. 91).
The immediate derivatives of 61 H. cymobrathys are three species of the Colombian and
Venezuelan Andes (see Part 8). Of these the Colombian H. chamaemyrtus is the sister-species of
the H. gnidioides — H. gentianoides group, which radiates from the Honduras Republic: south to
Panama (H. gnidioides), north to north-eastern U.S.A. (H. gentianoides), and east to the
Greater Antilles (H. diosmoides group). On the other hand, there is a wide geographical
disjunction between the Venezuelan H. pseudocaracasanum and its sister-group in the south-
eastern U.S.A. (H. denticulatum et aff.).
(b) Species with leaves persistent or deciduous near the base (Spp. 3-23)
In this group (Fig. 3 clade (c)) the leaves are rarely broadest above the middle, and so the
leaf-venation is parallel to pinnate or 1-nerved, not flabellate. All species have leaves that are
either wholly persistent or deciduous at the apparent base. They form two subgroups: clade (1)
with leaves that are free (Spp. 3-9); clade (ii) with leaves that are united by an interfoliar ridge or
rarely more intimately (Spp. 10-23).
Subgroup (i) radiates from 3 H. phellos of north-eastern Colombia and adjacent Venezuela.
The variation in this species can be described; but it has not been possible to express this
variation in a formal classification, apart from the recognition of two derivative subspecies.
Subspecies H. phellos has therefore had to be differentiated plesiomorphically (i.e. by the
possession of primitive characters), viz. by having plane or recurved (not incurved or revolute)
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 (César/Norte de Santandér 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
half way (asin, say, H. elodes), and two pairs of fascicles are more or less completely united. The
petals are oblique to erect, sometimes making the flower pseudo-tubular, again as in H. elodes.
How the gap between southern Ecuador and south-eastern Brazil was traversed is not clear;
quite possibly it was by ancient long-distance dispersal. I have not yet discovered any similar
disjunctions, but the morphocline southward along the Andes to Ecuador suggests that a
vicariant interpretation is unlikely to be correct.
An early offshoot of clade (ii) comprises two isolated species (clade ix) in which the relatively
large herbaceous leaves fade and droop while on the stem. 10 H. simonsii is confined to the
northern part of the Sierra de Santa Marta (Magdalena), whilst 11 H. papillosum has a restricted
distribution in Boyaca. In the remainder of the clade (ii) the leaves are smaller and either
broad-based and deflexed (clade xi) or narrow-based and spreading only (clade xii). At the
extremes of clade (xi) the leaves become too narrow to droop and so remain spreading (15 H.
myricariifolium, 18 H. loxense in part). The species at the base of these clades (13 H. thuyoides)
has a vicariad that is scarcely more than subspecifically distinct (12 H. lycopodioides), in which
the leaves have become coriaceous. H. thuyoides itself, from central Colombia (Cundinamarca),
is variable, with the broader-leaved (typical) form near clade (xi) and the narrower-leaved form
(‘genistoides’) at the base of clade (xii). Clade (xii), in turn, divides into two distinct subclades:
in one (xiv), the areas of 22 H. hartwegii and 23 H. maguire? represent a major disjunction to
southern Ecuador, whereas in the other (xiii), H. thuyoides is near the origin of two trends
initially in opposite directions, respectively northwards to Boyaca (19 H. sabiniforme) and
southward to northern Ecuador (20 H. laricifolium). As has already been mentioned, H.
laricifolium shows continuous variation southwards to central Peru and northwards to western
Venezuela; but the morphologically reduced population further north, in the Sierra Nevada de
Santa Marta (Magdalena) is sufficiently distinct to warrant specific recognition (21 H. martense).
(c) Species with leaves with sheathing leaf-bases or marginal secretions (Spp. 24-26, 61)
In contrast to the early offshoot (f) in Boyaca in which the oblong to elliptic leaves of the basic
species have parallel venation (61 H. cymobrathys), nearly all early members of clade (d) (Fig.
3) have leaves broader above the middle with flabellate venation. A partial exception is 24 H.
magniflorum, in which the Boyaca population has parallel-veined leaves, but the more northern
one (straddling the Colombia- Venezuela border) has flabellate venation. H. magniflorum is also
exceptional in that the leaves are relatively thin and droop when fading, instead of breaking off
above the base like those of all the other species. It forms the base of one sub-branch of Branch
(e) (clade xv), in which the broad, flabellate-veined leaves have sticky marginal leaf secretions,
capitate stigmas, and usually a cymose inflorescence. 25 H. gleasonii occurs in the border area of
Colombia/Venezuela, whereas 26 H. mexicanum has its primitive form in Boyaca and Cundina-
marca and differentiated back northwards into the area of 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 H. pimeleoides, which is found in the Cerro de Oroque (César/Norte de
Santandér), 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 H. costaricense, is confined to
the South American Cordillera (Norte de Santandér to Boyaca).
In the relict basic species of clade (xxiii), 40 H. bolivaricum (Colombia: Bolivar), twisting
leaves are combined with a condensed synflorescence and capitate stigmas, all characters that
are found in the variable 41 H. juniperinum (= H. brathys). In the latter species, which is
distributed from western Venezuela (Trujillo) to southern Colombia (Putumayo), there is a
trend towards an increase in the number of styles and placentae from 3 to 4 and then to 5. The
population from Cauca shows another trend, among 3—4-styled forms, from tall erect shrubs to
smaller shrubs with decumbent stems. This trend is continued back north-eastward in 42 H.
prostratum, a dwarf shrub or shrublet from the northern half of the Colombian Cordillera
Oriental with stems decumbent to prostrate.
The remaining derivatives of 32 H. magdalenicum are all related to more advanced forms of
that species (32’ in Fig. 3). Apart from 43 H. parallelum they form two clades: (xxvi) of which
the members have polished, acicular leaves, rounded-incurved in section and twisting when
large enough, and (xxvii) in which the species have leaves that are dull to metallic and flat to
incurved but not usually rounded in section and become recurved not twisted. Clade (xxvi)
comprises three geographically separate taxa that I have treated as subspecies of 44 H.
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: Mérida and Colombia: Boyaca) and differs from subsp.
marahuacanum in size and condensation of parts. In comparison with H. marahuacanum, the
species of the H. strictum group (Spp. 45—48) are south-western in distribution, being confined
to the area from Cundinamarca and Meta (Colombia) to western Venezuela. The relatively
broad-leaved 45 H. lancifolium, from the Colombia-Venezuela border area, has a dwarf,
high-altitude derivative, 46 H. horizontale. on the Colombian side of the frontier and a southern
derivative complex (47 H. tetrastichum, 48 H. strictum) with H. tetrastichum extending from
Boyaca northward into Venezuela and H. strictum occurring farther south.
(f) Species with pseudopetiolate leaves deciduous above the base but not incurved-cucullate (Spp.
31 p.p., 49-60)
The above heading is not strictly accurate in that a few of the advanced members of this group
(clade xix) have developed a somewhat cucullate apex. But their leaves are linear and would not
be confused with those of species in clade (xvi).
The species with pseudopetiolate leaves and their derivatives in clade (xix) (Spp. 49-60)
would appear to have evolved relatively early in the history of sect. Brathys, as they have spread
to almost the furthest limits of the shrubby part of the section (Hispaniola and Costa Rica to
Bolivia). The basic species of this clade, 49 H. jaramilloi, has a disjunct distribution in northern
Colombia (Cerro de Oroque, Sierra de Santa Marta) and Costa Rica; and an early species in the
eastern subclade (xxviii), 50 H. cardonae, has a similar but wider distribution, in Costa Rica and
the Colombia/Venezuela border region south to Boyaca. The rest of the species of this clade
show a geographical replacement pattern eastward: 51 H. caracasanum in the mountains of
northern Venezuela, with 51b subsp. turumiquirense at their extremity in Sucre, and the
remainder (Spp. 52-54) in Hispaniola (see p. 9).
The second (westward) derivative clade of 49 H. jaramillo, clade (xxix), is based on 55 H.
ruscoides, from south Colombia and north Ecuador, which gave rise southward to clade (xxx), a
series of sessile-leaved species (56 H. llanganaticum in central Ecuador, 57 H. struthiolifolium in
Peru, and 58 H. andinum in south Peru and Bolivia) with acute leaves and (at least initially) long
styles with relatively narrow stigmas. In another direction, evolution resulted in two species with
obtuse, cucullate leaves and short styles with relatively broad stigmas. Of these, 59 H. lancioides
comprises a southern subspecies with a pseudo-dichotomous synflorescence (59a subsp. /an-
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. Hypéric.: 38, 58 (1821).*
Small trees, shrubs, shrublets or wiry herbs up to 6 m tall, evergreen, glabrous or very rarely with
simple hairs, without dark glands; branching below inflorescence lateral. Stems 4(6)-lined and +
compressed (ancipitous) when young, sooner or later becoming terete, eglandular; cortex
exfoliating in strips or scales or irregularly; bark fissured, smooth. Leaves opposite, decussate,
sessile to shortly petiolate, free or + deeply united to form interfoliar ridge or very rarely bases
of lamina proper united, base sometimes sheathing, deciduous at base or (more usually) above
base or persistent and (in woody species) either fading or shed with cortex; lamina entire, with
venation parallel or flabellate to pinnate or 1-veined, open or closed, with tertiary venation +
laxly reticulate; laminar glands punctiform; marginal gland dots dense, often very small; ventral
* Including Sarothra L. (1753), Martia sensu Sprengel (1826) pro parte, Hypericum sect. Brathys subsect. Multista-
mineum R. Keller (1893). Hypericum sect. Brathys subsect. Spachium R. Keller (1893), Sanidophyllum Small (1924),
Sarothra sect. Spachium (R. Keller) Y. Kimura (1951), Sarothra sect. Spachium series Eusarothra Y. Kimura (1951),
see Part 1 (Robson, 1977: 338); also Hypericum sect. Sarothra (L.) Reichardt in Martius, Fl. Brasil. 12(1): 186 (1878).
THE GENUS HYPERICUM L. 13
resin glands absent. Inflorescence 1-flowered, with branching pseudo-dichotomous (usually
from uppermost node) or 2—15-flowered and then with branching (within inflorescence)
dichasial/monochasial from uppermost node, often with subsidiary branches from lower nodes;
bracts and bracteoles foliar or reduced or sometimes transitional in form to sepals. Flowers
stellate or occasionally obconic, homostylous. Sepals 5, free, persistent, erect in fruit, with
margin entire; veins 3—11(—19); laminar glands linear to punctiform; marginal, submarginal and
inframarginal glands absent. Petals 5, persistent, with apiculus subterminal to obsolete, acute to
acuminate or obtuse; margin entire; marginal glands absent; laminar glands linear to punc-
tiform. Stamen fascicles basically 5, very rarely distinguishable individually, usually forming a
continuous narrow ring of 5—250 stamens (i.e. 1-50 per fascicle) or very rarely fascicles + united
2 + 2 + 1, persistent; filaments very shortly united or very rarely united above midway; anthers
yellow to orange, gland amber; pollen type VIII. Ovary with 3-S parietal placentae, ©-ovulate;
styles 3-5, free with bases contiguous; stigmas small to broadly capitate or clavate. Capsule
3—5-valved, coriaceous to chartaceous, with valves obscurely narrowly longitudinally vittate,
occasionally with 1-3 flattish vesicles. Seeds narrowly cylindric, shallowly carinate or not,
without apical expansion; testa scalariform-reticulate to ribbed-scalariform.
BASIC CHROMOSOME NUMBER (x): 12; ploidy 1, 2, 4.
Hasirat: 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 subpéramo
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, Chimantaé, 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
INTETIONUARTIOBES oonicc re cednce poeta tes ety Conn pubenatetels drat seebe wv antedee sewtidetepmbunnceNs 2
Leaves persistent (falling with cortex) or deciduous above base; stem articulate and/or with
INCETCOMAE TIQGES ower os adeseacan ei ee wa adultes acco s ec uues ra csckb cpuv'uwede nue boos hanes naneadE os a
2(1) Leaves narrowly elliptic, acute, 16-30 mm long, chartaceous to subcoriaceous; sepals
PDD WAIN ONG oh con vcs. © baie cee Sodus cuca eRieeh ep dean RRM eRCe get 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)......................:000605 3
3(2) Leaves narrowly elliptic to narrowly oblong, usually thinly coriaceous, spreading, often
OCCUNVIIE Gre ave lane tet ou canctyeldraknce cote aoaanaeaes 2a. styphelioides subsp. clarense(p. 21)
Leaves oblanceolate to narrowly obovate, + thickly coriaceous, + densely imbricate,
SUL Ne ae vac ilar Pan eg sues sete snes He ek a TN Puy sur da vautws inverters cornea ths hewteu kage ated 4
4(3) | Leaves with 7-11 basal veins, glands not prominent
2b. styphelioides subsp. styphelioides(p. 21)
Leaves with 5—7 basal veins, glands prominent......... 2c. styphelioides subsp. moaense(p. 22)
5(1) Leaves persistent or deciduous immediately above base of free part; pseudo-dichotomous
TANCES OLE AUSENU OL SAIS ricer ennai enresccawe esheets eam cusesemnpenterstesanrzo steed 6
Leaves deciduous above base, i.e. leaving base protruding from stem; pseudo-
chehotomous branches O1ten PregOMINANl 6.24565 365 ser. ou,se oases cess aiwnceeedn esses sn punssoscevene 38
6(5) Leaf bases either (i) sheathing or (ii) amplexicaul with stem fragile and articulated, the
Hmermodes and leaves falling 25 ONG 6 ...o< lociccssccnts ae saasducsGGarcacvesserietec ioe noudeeesiseennanee 7
Leaf bases free to perfoliate or shallowly cupped; internodes and leaves not as above ........ 8
14
7(6)
8(6)
9(8)
10(9)
11(10)
12(11)
13(10)
14(13)
15(14)
16(15)
17(16)
18(15)
19(13)
20(19)
21(20)
22(8)
23(22)
NORMAN K. B. ROBSON
Leaf bases sheathing; leaves 10-18 mm long, withering; flowers 25—40 mm in diam.
24. magniflorum(p.
Leaf bases amplexicaul; leaves up to 2 mm long, falling with internodes; flowers 5—7 mm in
GUDARR Da siete cas otcae soos horace eectea conc Stay ai Sele ei-ea tan center war eepeaaostaars 54. millefolium(p.
Leaves free, the base sometimes indurated or fused with the stem .................ccececeeeeeeee es
Leaves united by interfoliar ridge or lamina prope? ........0...:i::0sc0esssesseeconennntnarssoroesees
Leaves becoming deflexed, marcescent; stem internodes without or almost without corky
CMEFQENCES 5.5 hone cwssldge eh sseapedealantaeees 3a. phellos subsp. phellos (i) ‘marcescens’ (p.
Leaves becoming deflexed to appressed but not marcescent; stem internodes with or
WItHOULCOLKY CMNOrBENICES co 2 ys voc2 bcc uy cp avon apy recone a4 sien on open pte guneane face “bees naseeet scene
Leaves revolute, lower surface obscured, bases forming + woody projections; stem
PLETTIOGES SINGER. acres eG dae pene saske 0 dacivae cc adeeb Genes Ceslerneurs as tesewasesesacioneveseserwcryss
Leaves recurved to incurved, lower surface always visible; stem internodes usually with
CORKY CMICTOCTICER SG 55.5 non easel esstecerene Wen ducen t eege we vaste ae Ses Paw tense are sete es etak comune:
Leaves deciduous, 4-10 mm long, elliptic-oblong to linear; sepals ovate to lanceolate;
SHU EO OEY tal nary etek Nera AE asides rad Aoanrap nat ae heehee he ounnaas nian aeeate ane 4. garciae(p.
Leaves persistent, (8)10—40 mm long, linear; sepals narrowly lanceolate to linear or
oblong; dwarf shrub to 0-8 m tall or suffrutex Or Berd 2.556. 265035cc0ssessccesnrenssecesersusso nce
Flowers solitary, inflorescence-branching pseudo-dichotomous; flowers stellate; stamens
not fasciculate;.dwart Shrub: &..,.:..4¢ 14. 5.5135 as ed jacaesagsssateessades sscseshwmes 5. acostanum(p.
Flowers 1-25, inflorescence-branching monochasial; flowers obconic to pseudotubular
stamens 3(5)-fasciculate; suffrutex or perennial herb ...................:seeeeeeeees 6. piriai(p.
LOAVES With :o bdSal VEINS. SUDCOFIACEOUS sass ca25 235 ory tse ou Buy ndansak onensscaseneneateatatsenagertnanes
Leaves with 1 basal vein (venation wholly pinnate), chartaceOuS................scceeseeeereeeeeees
Leaves recurved before falling ................... 3a. phellos subsp. phellos (ii) ‘angustum’ (p.
Leaves spreading to appressed before fallin. jive vcrce ce ciese ts: avec Waves rises don sentese soooxeene ts
PG aVies CEGIGMOUS cet caetias cas sas ths saa Aba BE cau esniedaivaldcaeeleeecatenees nneseae cares eeme seas
PE GAVGS POISISTO BG sce co nattars Adi oks dace sutsnicatms candugealeaupdyerecleben tirchesemaesnub asa ueaoenn opts
Leaves spreading, not or scarcely becoming appressed, elliptic to oblanceolate
3a. phellos subsp. phellos (iii) ‘patens’ (p.
Leaves becoming or remaining appressed, oblanceolate to narrowly oblong................++++
Leaves spreading, then appressed, before falling
3a. phellos subsp. phellos (iv) ‘phellos’ (p.
Leaves remaining appressed, tetrastichous ............... 3b. phellos subsp. oroqueanum(p.
Leaves with 3 unbranched basal veins; flowers 25-30 mm in diam.; styles 9-10 mm long,
BOW AS ONG AS OVALY ces coun <ace seman ates n do teases fn dea nde taninsecuaeda ce vgAwonewes 7. irazuense(p.
Leaves with at least midrib branched; flowers 15—20 mm in diam.; styles S—9 mm long, 2—4
MOS TOME BS OVADY essa eatn svn asso ve dele oie aed vanes waemda t erauNe we eae ap 8. stenopetalum(p.
Leaves deciduous or subpersistent; midnb branched’: ;. i: 2. scission sco igs oes sb on va cecens dase ses
Weenves persistent. 1 -merved 55 cc5 3 foc rk casos te babes sdues seseanraoeinar ane nsas ee 9. carinosum(p.
Leaves with margins plane or recurved, spreading or becoming ascending, deciduous........
Leaves with margin incurved, closely appressed, subpersistent
3c. phellos subsp. platyphyllum(p.
Leaves broadly elliptic to obovate, not becoming ascending
3a. phellos subsp. phellos (v) ‘diversicaule’ (p.
Leaves narrowly elliptic to oblong, spreading or becoming ascending
3a. phellos subsp. phellos (vi) ‘tamanum’ (p.
Inflorescence-branching cymose; leaves always broadest at or below middle, rarely
petiolate, with margin plane or incurved, not minute and scale-like ..............:sseeeeeeeeee
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
Sale TG aides shes eet faces MRR TE ARES nae eons an etude vcepenie yeaa Cons Taas cae pa vsatad decane cian anaed
Leaves plane, oblong, with 3—7 basal veins becoming deflexed when withering; sepals 6-11
POI 5 o.oo sissn c see ueae x oad tess wha Sho Saas canaeanpawates consee ees eie eres neciasiaiss praca aes
26)
27)
28)
14
19
25)
15
16
18
25)
17
25)
25)
30)
31)
20
33)
21
25)
25)
25)
24(23)
25(23)
26(25)
27(26)
28(26)
29(28)
30(28)
31(30)
32(25)
33(32)
34(33)
35(32)
36(35)
37(22)
38(5)
THE GENUS HYPERICUM L.
Leaves incurved or involute or conduplicate, or if plane then broadly ovate or 1-nerved,
becoming deflexéed or not: sepals 2-6(—7) mm JONG. ; 20... ci ccceen cs cae asdessnssdsaccseives asees
Leaves smooth, + lustrous, the pairs very shortly united; stigmas narrow ..._ 10. simonsii(p.
Leaves papillose, dull, the pairs united to form a cup-shaped sheath; stigmas capitate
11. papillosum(p.
Leaves united only by interfoliar ridge, not becoming deflexed (except sometimes 12.
LI COM OGINGRS) an Fa ceattrg ceeds tReet SEEN Nitec ob ae od Mes aR aA De ER EE
Leaves with bases of lamina united, often becoming deflexed....................cccccccececeee eee es
Leaves with midrib proximally prominent or wholly level with lamina; lamina condupli-
Cate “APEX UISUAlLY ACHININ ATC (ODUUISE .i.coscapeucn, owners. Was sPaceeu tuk 1h ab wien Hae te cones esse eee
Leaves with midrib proximally impressed; lamina incurved, apex usually obtuse to
POUNGCO reas cage hese c maar cb caren phen ete MR bare aa sae NIAC Salt see OTL oe pclae eT
Leaves coriaceous, markedly tetrastichous, apex acuminate to acute, margin plane
12. lycopodioides(p.
Leaves chartaceous, weakly or not tetrastichous, apex acute to rounded, margin usually
AINE so hvc ce sit 2 oy oh Vara tay ucla ag a nda epee eae oe ne crevice Soaneneaee 13. thuyoides(p.
Leaves sessile, elliptic-oblong to narrowly oblong, base cuneate to parallel .....................
Leaves shortly petiolate or base angustate, if base narrowly cuneate, then lamina linear to
PCICUNAL 7. ot fen er Sede rs Heda Nc deer enact eaeed toe Nai air Tea ata ees aS oa eae See
Leaves 5—7 mm long, elliptic or oblong to obovate; sepals 1-8—2-3 mm wide, apex plane
22. hartwegii(p.
Leaves 2—4 mm long, narrowly elliptic or narrowly oblong to linear; sepals 0-7—1-7 mm
Wide; apex inCirved-CuCUllate.s 15 <.wa-oy. eae ook ae oeec ws mec ecsangeveueeeres 23. maguirei(p.
Leaves subpapillose beneath, base cuneate to pseudopetiolate; sepals obtuse to rounded
19. sabiniforme(p.
Leaves smooth beneath, base narrowly cuneate to angustate or parallel-sided; sepals acute
LO ODUUSE 5 EF A cay eee edak cutsecs puavetinae so wusleasen Bote ates tea ek UU Laa Ra senac aie osigs Date ee ee
Plant a bushy or spreading shrub or small tree (0-1)0-3—6 m tall; leaves with glands not or
Scarcely impressed: sepals ustdlly DlANG< pcr1c hee rci set nates vice ncveecese: 20. laricifolium(p.
Plant a pulviniform shrub 0-08—0-16 m tall; leaves with glands impressed; sepals markedly
BUDO EG yarn essa ede can ec vas nats seen Hiea te sae de amob tote meartd aces duane eee 21. martense(p.
Flowers 18—40 mm in diam.; leaf base rounded to cordate-amplexicaul ......................0068
Flowers 6—12mmin diam. Sléaf base parallel to truncate: vcccca.cc.ses-2- sonst oeten esses tress Poneetee
Stem internodes smooth; leaves plane or slightly saccate, (2—)3—7 mm wide, margin not or
SCAICELY NYGNING San. cte canes ome aaa dee panic epee y eaemeaete 16. cuatrecasii(p.
Stem internodes with corky emergences, at least below upper leaves; leaves saccate or
incurved-cucullate, 0-7—3-5 mm wide, margin markedly hyaline........................c0eeees
Leaves becoming reflexed, saccate; sepals 5—8 mm wide....................... 14. goyanesii(p.
Leaves spreading only, incurved-cucullate; sepals 4-5 mm wide..... 15. myricariifolium (p.
Leaves becoming deflexed, broadly ovate to triangular-ovate or (lower) oblong, basal
ETA SO ees ches ae dee cerca ce co tebe Staal wale airs ark tors apton ca pateetes 17. quitense(p.
Leaves at most spreading only, triangular-ovate or oblong to linear, basal veins 1(3—5) (18.
WIROMRO) Sica a8 einen b= Dc as Se ot ste ew netdai epsci= pase an tae te Sree es eee EP e en meena areenu mates
Leaves broadly triangular-ovate to lanceolate, usually imbricate, base truncate to broadly
CUNERLE 2 .criussscesg suc cecaut arneat Con eman ceed up esds tent res ens 18a. loxense subsp. aequatoriale (p.
Leaves narrowly oblong or narrowly elliptic to oblanceolate or linear, usually spreading,
base narrowly cuneate to parallel a. is cccssevassstuesessews 18b. loxense subsp. loxense (p.
Leaves with parallel venation (5—7-veined); flowers 3-12 in condensed cyme; shrub
34)
34)
26
32
27
28
36)
37)
29
30
51)
52)
45)
31
47)
51)
33
a)
41)
34
38)
40)
41)
36
44)
45)
61. cymobrathys(p. 103)
Leaves with pinnate venation or 1-nerved; flowers © in + lax cymes; suffrutices or herbs
[62-80. See Part 8]
Leaf venation parallel (S—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
ERGO Ie ZOSIMEICIILOIE) 1 foe oc cee tae CT Ass sss Oc Seas g sens hes ye Siack pepe ores e rien teeter nees
39
16
39(38)
40(39)
41(40)
42(40)
43(42)
44(43)
45(39)
46(45)
47(46)
48(47)
49(48)
50(49)
51(50)
52(50)
53(48)
54(46)
NORMAN K. B. ROBSON
Leaves and usually sepals + markedly incurved-cucullate; leaves distally broadened, not
GINO AT es cae st ete Tp mt tan se cee cette pean cee sanz cumah xwinat aeth gies taaeds ear tesseueeoeees
Leaves plane to incurved; leaf and sepal apices not cucullate or, if so, then lamina narrowly
elliptic ornarrowly oblong tolineat y s.0. wen sae concen ne cone sen extevan nests angoaameVedou ae asticenes des
Leaves with margin viscid, usually obovate; flowers often in dense cymes; stigmas +
CADILRES se ioc he cote Se tor scenes cn eceh Non eusnmae aD Oo Coie cad ace « etn eve eu eee Tee eeu Ona r an
Leaves with margin not viscid, broadly elliptic to obovate-spathulate; flowers solitary;
stigmas narrow or, if capitate then leaves oblanceolate .................ccseeeeeeceeeeeeeeeeneeeees
Flowers solitary, 20—40 mm in diam. , sometimes terminating clustered short shoots; leaves
7—10mmiong, always coriaceous; habiterect 1 ....c2scwencecss cess st hans teas 25. gleasonii(p.
Flowers in 3—13-flowered cymes or, if solitary, less than 20 mm in diam.; leaves up to 17
mm long, sometimes chartaceous and then habit decumbent and rooting
26. mexicanum(p.
Leaves elliptic-ovate to elliptic-lanceolate, 3-7 mm broad venation flabellate; flowers
25a Tne WANN eee oe eae os ee reee Net re te ee Wes er ean Seer L apiece 27. stuebelii(p.
Leaves oblanceolate to narrowly oblong, 0-7-3 mm broad, venation pinnate or 1-nerved;
flowers6—2)5 mm in diam.( petals unknown in 28. prietoi) )i.0 0205 joceca cs 1085s sea canece ctansesee
Sepals acuminate; leaves persistently imbricate-tetrastichous, markedly laterally com-
pressed distally stigmas Capitate: coo cer. tec we ect teeter oaeeern eo eanes 28. prietoi(p.
Sepals acute; leaves eventually outcurving, not laterally compressed distally or, if so (29.
cassiopiformein part) then'stigmas Narrow 9 c:o.,c05-25300 1, so scene Deucteatebicges steve eeatecass
Petals c.14 mm long; styles longer than ovary; stigmas narrow; leaves oblanceolate (1: b =
CDS) Docc dnd Hive te ees ee pn Rec egetaa hea Jane BE ate oan ee kins s PORE RELL 29. cassiopiforme(p.
Petals S—9 mm long; styles 0-6-1 X as long as ovary; stigmas capitate; leaves narrowly
oblanceolate to linear-elliptic (1: b = 4-11)... eee eee eee 30. decandrum(p.
Leaves with venation flabellate, some laterals free or almost free from the base; styles stout
with stignia usually capitate.) cacti ee ised ste ee cctees cesmeosaee cee: 31. pimeleoides(p.
Leaves with venation pinnate, all laterals originating well above the base; styles and
StIDIMAS VATIOUS i525, eiienevet en tee ead ovis awed eee save eaea eaten teubeesone aeeR enone ee
Leaves petiolate or abruptly narrowed at the base; stem sometimes apparently articulated
Leaves sessile, gradually narrowed or parallel-sided at the base; stem not articulated ........
Styles 6-7 mm long; flowers 25—30 mm in diam.; leaves often spreading abruptly above
aro (6) | oA RRS ROE CB ODT, Cena TSP iE rr AO CR a Re A Tee 49. jaramilloi(p.
Styles 0-8-5 mm long; flowers up to 25 mm in diam.; leaves outcurving or spreading from
ING DASE eos. cris cisaiy anion oe tscn ainda es dace casts tere einen Mo naxaenesG ac wes noun iandeosaaeenotenets
Leaves with midrib prominent or level with lamina beneath, glands usually visible beneath
Leaves with midrib + impressed beneath, glands not visible beneath .....................:2:0000
Stigmas + broadly capitate; leaves glaucous, dull..................cceeeeeee eens 50. cardonae(p.
Stigmas clavate or small; leaves not glaucous, + lustrous: 222.25 2icsekc sch 4.hakcesendevsevesevseties
Leaves with marginsrecurved(51, Caracasamunil) 56 (ccs cisco 50505205 sect ccgtseuavsraaresendeosteneses
Leaves withmaroin or whole lamina incurvedss....22:. foses-2cnpars ewiunsninanedyseacesicednssieowinas
Leaves 9 X 4-5 mm or larger; petals c.1-3 x as long as sepals; styles c.1-3 x as long as ovary
51a. caracasanum subsp. caracasanum(p.
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
QV AUN 5 ao earns nescrseohis amine camintan ee tunes 51b. caracasanum subsp. turumiquirense(p.
Leaf lamina elliptic, 2-4 mm wide, plane with margin incurved; styles 5-6 mm long
52. ekmanii(p.
Leaf lamina narrowly elliptic to linear, 0-5-2 mm wide, incurved to canaliculate; styles
Pe MME OND see tree tensed athe spears anaes ee pareenae eee 53. pycnophyllum(p.
Leaves plane or with margin subincrassate, narrowly ovate to narrowly oblong-elliptic;
sepals 1-5—4 mm wide; styles 2-5—4 mm long....................:eceeeeeeeeeees 55. ruscoides(p.
Leaves incurved to canaliculate, very narrowly lanceolate to (usually) linear; sepals
0-5—1-5 mm wide; styles 1—-2(—3) mm long............. 59a. lancioides subsp. lancioides(p.
Leaf and sepal apices + incurved-cucullate; leaf lamina narrowly oblong or narrowly
elliptic to linear: arrowed Alte Dase <.5.6es sa vecdenens cts sctncpsnysasancconsssaversccsaesunspsneenen
55(54)
56(55)
57(56)
58(55)
59(58)
60(54)
61(60)
62(61)
63(62)
64(63)
65(64)
66(65)
67(66)
68(66)
69(65)
THE GENUS HYPERICUM L. 1
Leaf and sepal apices not incurved or cucullate or, if so, then leaves broadened at the base 60
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 PTI DDE eis onsen cers dena cavechnvvnwadenenainncn eines 58
Stems erect or rarely decumbent; leaves 7-15 mm long; sepals 5—8 mm long; styles 3(4),
B—2(—3) Hany lon (D9 TAICIOND OS) oie ssc de vd oocg ons ee ane ees ae sage ota es «ee OE eR oe 57
Stems prostrate or ascending; leaves 4-7 mm long; sepals 3—4 mm long; styles (3)4—5,
Ose PIM LONG 4 ork cas ake eee ect Be as cae eee eet eau eea ke ide does 60. selaginella(p. 102)
Flowers terminal, solitary, inflorescence-branching pseudo-dichotomous; habit bushy or
CAPS DIROSE coc Jas dna tea Sy uese ae ance tinted asc ea sets 59a. lancioides subsp. lancioides(p. 99)
Flowers terminal and lateral in dense racemiform heads; habit bushy or
SUCRE SANG o-oo co derdeys pores tes srecrsmasetee cael s ss 59b. lancioides subsp. congestiflorum(p. 101)
Styles 3—4-5 mm long, 1-3—1-5 X as long as ovary; stigmas not or scarcely capitate; leaves
d:5—4:mim wide. margin not or scarcely valine 56, 02: 623 sicsds via nessangtariswaaueiotwaeccoas encee 59
Styles 1-5—2-5 mm long, 0-6—0-7 Xx as long as ovary; stigmas broadly capitate; leaves
0-8—1-5 mm wide, margin narrowly but distinctly hyaline.................... 58. andinum(p. 97)
Sepals oblong-oblanceolate, not ribbed; leaves plane or with margin slightly incurved, 6-9
mm long, markedly tetrastichous, glaucous ....................eceeeeeees 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. struthiolifolium(p. 96)
Styles 6-7-5 mm long; flowers 20-30 mm in diam.; leaves with midrib branched, lamina
elliptic to oblanceolate, plane or conduplicate,epidermissmooth 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 ...................06.05 61
Leaves markedly tetrastichous, narrowly elliptic, papillose above; branches very strict,
pataliel,withinternodés 1—1-5 mm long ane eo ees gs cs pne ee eas 43. parallelum(p. 76)
Leaves not tetrastichous or if so, then linear, smooth, and branches not as above .............. 62
Sepal margin green or, if hyaline, then inflorescence a racemiform synflorescence and
Styles shorter tian ovary with stipmas capitate css 16 i152 oes aR och es oes ea ete 63
Sepal margin hyaline; inflorescence and styles not as aDOVE ................ccececeeeeceeeeeceeeeeeen 72
Flowers solitary, not congested; styles 2-5—5 mm long, stigmas narrow...................0000000 64
Flowers in + congested heads or racemiform synflorescences; styles 1-3(4-5) mm long,
Stipmas + broadly capitate. xc gies: beassetecocat Auer tias. c's «1as vo Oran RN toe aoa 70
Leaves 2—3 mm wide, narrowly elliptic, slightly incurved..................... 33. valleanum(p. 64)
Leaves 0-4-2 mm wide, very narrowly oblanceolate to linear, markedly incurved to
CONGUDIICALE 50 tne, feccrccxcsun ghee roes hanes ie as sage ny th ls epee eh or enad as. GRA SRE Ben anda eece 65
Styles 2:5~3: Simm long; leaves lustrous beneath s2.715.55, ssessps en sate tases scares anon aaa eeeenteon 66
Styles l=-2imm long: leaves dull beneath icc. Sosges soos a eae fo eee vs co eae eee ee 69
Leaves becoming twisted, incurved but not carinate or conduplicate, margin very narrowly
hyaline; dorsally usually glandular when mature 2...csss..c-cc,0ces aces ore tieweste deckexCacseasnane 67
Leaves remaining erect or becoming recurved, incurved to conduplicate, often at least
partly carinate, margin broadly hyaline, dorsally eglandular when mature ................... 68
Sepals 7-11 mm long; flowers 20-30 mm in diam.; capsule c.5 mm long; leaves 6-12 x
OSU ET WA SEAIIEIIS OU 1a euwe tan coaie ec ose pre oreceia Ses teaser tente cs 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
SFBe ON SEAINETIS BU 00) oe ors cute tan canada ata sya toe > se vaTIAG Coates Catena 35. aciculare(p. 66)
Leaves imbricate at first but not markedly tetrastichous, recurving, 5-8 mm long; flowers
152 min in diam.; petals $=12 min long 7.32, .0250. cc cceun-esecevs onneo 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)
Flowers 15—20 mm in diam.; leaves 5—15 mm long, acute; stems woody, not rooting
38. costaricense(p. 70)
18
70(63)
71(70)
72(62)
73(72)
74(73)
75(72)
76(75)
77(76)
78(77)
NORMAN K. B. ROBSON
Flowers 6—8 mm in diam.; leaves 3-5 mm long, subacute to obtuse; stems wiry, basally
POORIELG os coarse oh ceo sa eee ea eee cat oy eras eta teres Fneeonniaee Fare ans 39. bryoides(p. 71)
Leaves very narrowly elliptic, 15 x 3mm or larger; sepals 9-11 mm long, coriaceous; styles
CoS IMLIOUS Sees seach ete ew ovce dn ens ee oy aan Yo caine ope nce Reiccsiog tees 40. bolivaricum(p. 72)
Leaves linear, up to 14 x 1-5 mm; sepals 2—7 mm long, chartaceous to membranous; styles
Eo hs 1110 (6) 1s Jan es ae Aneta af SPR Seances Sarre M AAP REE PU im ry re Me Envi dion tame man Sr L v1
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)
Leaves lustrous, margin markedly incurved, lamina twisting and sometimes recurving (44.
Pit Tg Li) te 111117 /) NASSP OSSSREREED SEE TT Grr tte REE Re CHT re ee ee roe E Pre etd dee BU sn ces v8)
Leaves dull or with metallic sheen, margin plane to incurved, lamina recurving only.......... 75
Stigmas small or slightly enlarged; ovary and capsule acute; inflorescence branching
Pscudo-dichotomolus OF TACCMMOTA can pat con dves sopnaete enone cece scwcunet keek Tix neeasaaineeee 74
Stigmas enlarged to capitate; ovary and capsule + rostrate, inflorescence branching
pseudo-dichotomous to lateral-congested 44c. marahuacanum subsp. chimantaicum(p. 80)
Styles 4-5 mm long; sepals 7-9 mm long; inflorescence branching pseudo-dichotomous
44a. marahuacanum subsp. marahuacanum(p. 79)
Styles 2-3(—4) mm long; sepals 4-6 mm long; inflorescence branching lateral, + racemi-
| (ay 911 War rare OM SY AP oe Taree eee er ae! 44b. marahuacanum subsp. strictissimum(p. 79)
Leaves plane or slightly incurved, oblong to oblong-linear or narrowly oblanceolate, the
tips NOtTEG WHEN YOUNG saree. re... race. 255. et eaaee sca Fa suanh Gen ¥ veer sores 45. lancifolium(p. 80)
Leaves markedly incurved to canaliculate, linear to linear-acicular, the tips usually red
Wile YOUN Bais sont eee epee te sates ea ented er ere teads ve eRaene ss deca eek tec netes Rogrolmiweah rr ceeih 76
Plant a prostrate shrublet with branches matted, rooting; leaves often secund, not
POET ASTICIOUS SS oe tirs eee ies eed sh ope ewed Came iesee ach oe estes 46. horizontale(p. 81)
Plant an erect shrub with branches strict, not rooting; leaves never secund, usually +
CEEEASTICHO US OC Rees ata eis de sages ees de Teen ee anaes poet tins venvaticts Muviseds 77
Leaves with metallic sheen, markedly tetrastichous, erecttosuberect 47. tetrastichum(p. 82)
Leaves dull, not tetrastichous (except sometimes in young parts or plants); usually
MULCUE VANS (ORs SeriChaIn As cat yy cote ee ees a sweeeone a artes sche Rn eee aes cates Codpaeea eee 78
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 S—7(8) mm
long; branches ascending, usually branched from the base
48b. strictum subsp. compactum(p. 85)
1. Hypericum terrae-firmae Sprague & Riley
321 (K!, holotype; GH!, isotype).
Adams in Rhodora 64: 236 (1962), non A. Richard (1845).
Icon: Fig. 4.
in Kew Bull. 1924: 12 (1924). Type: Belize, without precise locality, 1912? (fr), Peck
H. styphelioides sensu Standley & Williams in Fieldiana Bot. 24 (7): 51 (1961); P.
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, 7 = 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.
by
ae
SSN
/
N
Lf
ZL,
> : yi . N \
rN
Aix
Ll
N
N
=
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 = 2; c X 1; d,j x 2; e-g
x 3;h x 8;i x 15). (a—d, g, j) Hunt 66; (e, f, h, i) Sutton et al. 193.
pairs free; basal veins 7, subparallel-sided, with short lateral branches, tertiary
reticulation not visible; laminar glands dense, not prominent. Inflorescence 1-
flowered, with pseudo-dichotomous branches from 1—2 nodes below; pedicel absent
or very short; upper leaves transitional. Flowers 40—60 mm in diam., stellate. Sepals
14-22 x 3-7 mm, narrowly oblong, acute; veins 15, dichotomising and reticulate
distally, with midrib prominent; glands linear. Petals bright yellow, 20-30 x 7-12
mm, c. 1:3 X sepals, oblanceolate; apiculus acute; glands linear, mostly uninter-
rupted. Stamens c. 250, longest 9-11 mm long, c. 0-3—-0-4 x petals. Ovary (2-5—)3
20
NORMAN K. B. ROBSON
—4-5 x 2-2-5 mm, + narrowly ovoid; styles 5, 5—7 mm long, 1-4-3 x ovary, erect,
outcurved below apex; stigmas small. Capsule 8-10 x 5-7 mm, broadly ovoid,
shorter than sepals. Seeds c. 1 mm long, not or shallowly carinate; testa finely
scalariform-reticulate.
In open pine or oak-pine forest or pine savannah on granite, often near streams;
lowland to 550 m.
Belize (Cayo, Belize). Map 1.
BELIZE. Belize: Colonel English Pine Ridge, Belize-Cayo road, 20.1.1958 (fl),
Gentle 9515 (DUKE, F, NY, S, US); Burrellboom, c. 9 m, 22.11.1970 (fr), McDaniel
12853 (F). Cayo: Mountain Pine Ridge, San Miguel, 400 m, 2.vi.1979 (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 (H. revolutum subsp.
keniense (Schweinfurth) N. Robson), a subspecies of Sect. 1 Campylosporus.
2. Hypericum styphelioides A. Richard
in Sagra, Hist. Phys., Pol. Nat. Ile Cuba (Bot. — Pl. Vasc.): 237 (1845); Leon &
Alain, Fl. Cuba 3: 316, t. 141 (1953); Lippoldin Wiss Z. Friedr.-Schiller Univ. Jena
(Math.-Nat. R.) 19: 378, f.2. (1970). Type: Cuba, [Pinar del Rio, near San Diego
de los Banos], Sagras.n. (P, holotype).
Shrub 0-15—2 m tall, erect, with branches erect, strict, pseudo-dichotomous or
lateral. Stems orange-brown, 4-lined when young, soon terete, without epidermal
wrinkles, cortex exfoliating in strips or scales; internodes 2-6 mm long. Leaves
sessile, spreading from above the base to closely imbricate and tetrastichous,
deciduous at the base without fading; lamina 5—25 x 2—8 mm, narrowly obovate to
narrowly elliptic or narrowly oblong, plane or incurved, sometimes + concave, not
carinate, concolorous, lightly to densely glaucous, thinly to thickly coriaceous; apex
shortly acuminate to acute, base cuneate to scarcely angustate, not or slightly
sheathing, pairs free; basal veins 5—9(—11), subparallel to flabellate, branching
distally to form a lax reticulum, tertiary reticulum not visible; laminar glands dense,
+ prominent. Inflorescence 1(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—S0(—70) mm in
diam., stellate. Sepals 5-15 x 2-—S 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 H. styphelioides have been recognized. Subsp. clarense, the most
similar to H. terrae-firmae, has a disjunct distribution (central Cuba, western Cuba,
Isla de Pinos), whilst the others are respectively eastern (subsp. moaense) and
western (subsp. styphelioides).
THE GENUS HYPERICUM L.
aN
vA
Map 11. H. terrae-firmae @.2. H. 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.1911 (fl), Bro. Clemente 2884
(LS, holotype; NY!, isotype).
Hypericum incurvum Urban, Symb. Anitill. 9: 405 (1925); Le6n & 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.1920
(fl), Ekman 12072 (S!, holotype).
Shrub 0-15—0-6(-1) m tall. Leaves 10-18 x 2-5-5 mm, narrowly elliptic to narrowly
oblong, thinly to thickly coriaceous, not or lightly glaucous, plane to concave or
incurved, spreading or outcurving; apex acute; basal veins 5(7). Flowers 15-30 mm
in diam. Sepals oblong to lanceolate, acute or obtuse, 5-veined; glands prominent.
Petals 7-16 mm long. Styles 5, 3—5 mm long.
Savannahs, fields, and coastal sands, sometimes on weathered dolomite; lowland to
montane.
In Las Villas, western Pinar del Rio and Isla de Pinos.
CUBA. Pinar del Rio: N. of San Diego de los Banos, 13.iv.1900 (st), Palmer &
Riley 535 (NY, US); mountains near E] Guama, 6.iii.1900 (fr), Palmer & Riley 160
(NY, US). Las Villas: Lomas de Banao, i.1920 (fl), Lund 123 (NY); Trinidad Mts,
Buenos Aires, near La Sabana, 5.viii.1936, Smith, Hodgson & Gonzales 3381 (F,
NY, S, US). Isla de Pinos: Siguanea, 26.ii1.1916 (fl), N. L. Britton, E. G. Britton &
Wilson 14926 (F, NY, S, US); between Mine de Oro and Playa del Soldado,
12.1.1956, Killip 45389 (US).
Subsp. clarense forms a cline from eastern Las Villas (where the flowers are larger
and the leaves are broader and flat) via eastern Pinar del Rio to Isla de Pinos (where
the flowers are smaller and the leaves narrow with an incurved margin and out-
curving towards the apex).
2b. Hypericum styphelioides A. Richard subsp. styphelioides
Shrub 0-3-2 m tall. Leaves 8-25 x 4-8 mm, oblanceolate to narrowly obovate,
plane, + densely glaucous, + thickly coriaceous, spreading to densely imbricate;
apex shortly acuminate; basal veins 7—9(—11); glands not usually prominent. Flowers
15—50(—70) mm in diam. Sepals elliptic to obovate, acute or obtuse-apiculate to
obtuse, 9-17-veined. Petals 10-22 mm long. Styles 5, 3-5 mm long.
22
NORMAN K. B. ROBSON
In pine forest or savannah sand or in wet places; lowland.
In Isla de Pinos, Las Villas and Pinar del Rio.
CUBA. Pinar del Rio: El Sabalo, Finca Sabanalamar, near sea level, 22. xiii. 1937
(fl), Killip 32263 (NY, US); Herradura, 1904 (fl), van Hermann 562 (BM, F, K, NY,
P, US); near Laguna Alcatraz Grande, 16.xii.1911 (fl), Shafer 11016 (NY, US). Las
Villas: Banao Mts., Sabana de la Gloria, 29.vii.1918 (fl), Léon & Roca 7928 (NY).
Isla de Pinos: San Pedro and vicinity, 12.1i.—22.11.1916 (fl), N. L. Britton, E. G.
Britton & Wilson 14148 (F, NY, S, US); Los Indios, 4.ii.1953 (fl), Killip 42603 (NY,
LIS);
Subsp. styphelioides forms two incompletely correlated clines eastward, showing
reduction in size and increasing leaf-imbrication. The only specimen seen from Las
Villas (Léon & 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, Plano el Sierra de Moa, Bisse 11676 (JE, holotype; HAJB,
isotype).
Shrub 0-6-1-5 m tall. Leaves 7—11(—16) X 2-6 mm, obovate (or rarely oblanceolate)
to narrowly elliptic, thickly coriaceous; slightly glaucous, + cucullate or incurved,
usually + densely imbricate; apex apiculate-obtuse to acute; basal veins 5—7; glands
prominent. Flowers 15-20 mm in diam. Sepals lanceolate, acute, 7-veined. Petals
8-12 mm long. Styles 5(4), c. 3 mm long.
In pine woods and open, dry or marshy habitats, on laterite or serpentine soils;
0-800 m.
Confined to Oriente.
CUBA. Oriente: Minas de Iberia, Taco Bay, in mountains, c. 800 m, 7—8.xii.1914
(fl), Ekman 3787 (F, K, MICH, NY, S); Moa, near airfield, vii.1941 (fl), Howard
6020 (BM, NY, P, S, US); Maravi (near Barancoa), 25.xii.1914 (st), Ekman 4020
(NY,.5; 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 H. phellos. On the other main division of H. phellos
subsp. phellos, (v) ‘diversicaule’ is related to 9 H. carinosum, which has smaller,
THE GENUS HYPERICUM L.
1-nerved leaves and smaller flowers, whereas (vi) ‘tamanum’, which differs from
‘diversicaule’ only in its relatively broader and more obtuse leaves and sepals and
more clearly differentiated lateral shoots, gives rise to a high-altitude condensed
form (3c subsp. platyphyllum) in which the leaves become incurved and remain
appressed until they are shed.
3. Hypericum phellos Gleason
in Bull. Torrey bot. Club 56: 106 (1929). Type: Colombia, Santandér, Paramo de la
Puentes, above La Baja, 3700-3800 m, 25.1.1927 (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 (1-)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 1(2—5)-flowered, terminal and on short lateral shoots, with extension
shoots usually from 2—3 nodes below; peduncle and pedicels 1-9 mm long; upper
leaves foliose or bracteose. Flowers 15—30 mm in diam., stellate. Sepals 4—9 x 1-5-3
mm, + narrowly oblong to lanceolate or elliptic or rarely oblanceolate, acute or
acuminate to obtuse or rarely rounded; veins 5—7, branched distally, midrib not or
scarcely prominent; glands mostly linear or up to half punctiform. Petals bright
yellow to orange-yellow, sometimes margined red, 8-15 x 4-8 mm, c. 2 X sepals,
oblanceolate to narrowly obovate; apiculus acute; glands all linear or distally
interrupted. Stamens c. 40-200, longest 5-8 mm long, c. 0-5—0-7 x petals. Ovary
2-3-5 X 1-5-2 mm, ovoid; styles 3, (5—)6—9 mm long, 2-3 X ovary, spreading,
incurved distally; stigmas small or subcapitate. Capsules 4-5-5 x 3-5-5 mm,
subglobose to globose, shorter than sepals. Seeds c. 1 mm long, ecarinate; testa finely
scalariform.
On scrubby slopes and in moist woods immediately below them; 2600-4200 m.
Colombia (Santandér, Norte de Santandér, César), 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
Santandér, Paramo de Tama, vicinity of La Cueva, 3100-3200 m, 27.x.1941 (fl),
Cuatrecasas, Schultes & Smith 12647 (COL, holotype; BM!, F!, U!, US!, iso-
types).
H. thymifolium Cuatrec. in sched., non Kunth.
H. tachirense Steyerm. in sched.
Shrub or small tree 0-3-4 m tall, with shoots rounded to pyramidal or narrowly
cylindric, not or + markedly differentiated into elongate main stems and short
laterals; internodes with slight or pronounced corky emergences. Leaves free,
23
24
NORMAN K. B. ROBSON
80
ia
Map 2_ 3. H. phellos (all subspecies) @; 10. H. simonsii A; 11. H. papillosum O.
spreading then deciduous directly or after decurving or after deflexing and withering;
lamina 5-5—16 x 1-4-7 mm, oblanceolate to elliptic or oblong, acute to acuminate,
margin recurved to plane. Sepals 4-9 x 2-3 mm, narrowly oblong or elliptic to
lanceolate or ovate-lanceolate. Petals 10-15 x 4-8 mm. Stamens c. 40-200. Styles
5—9 mm long.
Colombia (Norte de Santandér, Santandér), Venezuela (Tachira); 2500-3900 m.
COLOMBIA. Norte de Santandér: Paramo de Fontib6n, 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 Chitag4, 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).
Santandér: Péramo de Santurban entre Bucaramanga & Berlin, W. side, c. 3200 m,
3.i1.1960 (fl & fr), Barclay & Juajibioy 10422 (NY); Berlin to Picacho, 3058 m,
14.viii.1977 (fl), Renteria, Mantilla, Nino & Ortiz 555(3) (COL, MO); vicinity of La
Baja, 2700 m, 14—31.i.1927, Killip & Smith 18756 (GH, NY, US).
VENEZUELA. Tachira: Péramo de Batallén, near La Grita, 3100 m, 8.i11.1979
(fr), Kieft 104 (BM, U, VEN); SE. of Paramo de Tama, Pata de Judio, near the
Colombia-Venezuela frontier, 2900-3000 m, 20.i.1968 (fl), Steyermark & Dunster-
ville 101230 (NY, VEN).
The following is a summary of the variation of H. phellos subsp. phellos. The
variation is continuous, thus preventing the recognition of these variants as taxa. The
THE GENUS HYPERICUM L.
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 (Santandér: 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 Santandér: Paramo de Fontib6én, Mutis-
cua).
Variant 3a (ili) ‘patens’: Stems not differentiated, internodal corky emergences
evident; leaves deciduous when spreading, oblanceolate to elliptic, acute to obtuse,
margin recurved to plane (Santandér: Paramo de Santurban, Norte de Santandér:
Rio Chitaga4, Tachira: Paramo de Batall6n).
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 (Santandér: 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 Santandér: Péramo de Fontibén,
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 Santandér and Tachira: Paramo
de Tama, Santandér: Paramo de Almorzadero).
The type specimen of H. 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 Santandér y César, 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 Santandér/César); 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 Santandér, Paramo de
Romeral, 3800-4200 m, 30.i.1927 (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 4mm. Stamens c. 80. Styles 5-5-7 mm long.
25
26
SF
NORMAN K. B. ROBSON
Colombia (Norte de Santandér, Santandér), confined to the Paramos de Santurban
and de Romeral; 2700-4100 m.
COLOMBIA. Norte de Santandér: 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.1973 (veg), Cuatrecasas & Jaramillo
28751 (BM, COL, US). Santandér: 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, Santandér, Paramo del
Almorzadero, Peralonso, 3200-3300 m, 20.vii.1940 (fl), Cuatrecasas & Garcia-
Barriga 9935 (F!, holotype; NY!, US!, isotypes).
Hypericum chamaemyrtus sensu Gleason in Bull. Torrey 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.1944 (fl), Steyermark 57212 (F!, holotype; NY!, isotype).
Shrub 0-3-1-2(-3) m tall, erect, with branches strict or ascending, lateral or
occasionally pseudo-dichotomous. Stems orange-brown to blackish, 4-lined and
ancipitous when young, the principal lines broader, eventually terete, without
epidermal emergences, cortex exfoliating in strips; internodes 1-4 mm long. Leaves
sessile, outcurving or ascending and subimbricate, tetrastichous, deciduous at
suprabasal articulation; lamina 4-10 x 0-5—2-3 mm, elliptic-oblong or narrowly
lanceolate to linear, broadly revolute and often concealing lower surface apart from
midrib, not carinate, concolorous, lucent above, papillose beneath, not glaucous,
coriaceous; apex acute, base cuneate, not sheathing, pairs united to form a narrow
interfoliar ridge; basal vein unbranched; laminar glands dense, slightly prominent.
Inflorescence 1(2)-flowered, terminal and on short lateral shoots, occasionally with
pseudo-dichotomous branches; peduncle and pedicels 1-5-3 mm long, upwards
incrassate; upper leaves foliose. Flowers 17-20 mm in diam., stellate. Sepals 4-9 x
1-7-3 mm, lanceolate to ovate, acute to subacuminate; veins 5—7, obscure, not
visibly branching, midrib prominent; glands linear, distally punctiform. Petals bright
to deep yellow, sometimes tinged red, 8—13 x 3-6-5 mm, 1-5-2 X sepals, narrowly
obovate; apiculus acute; glands linear, interrupted distally. Stamens 40-70(?-100),
longest 5—8 mm long, c. 0-6—0-65 x petals. Ovary 1-5—2-5 x 1-2 mm, ovoid; styles
3(4), 3-5-7 mm long, 2:2—2-5(—3) X ovary, free, spreading-incurved; stigmas small
or slightly capitate. Capsule 4-5-5 x 3-3-5 mm, ovoid to subglobose, shorter than
sepals. Seeds c. 1:3 mm long, ecarinate; testa finely scalariform.
In paramo, usually on dry stony or sandy soil; 2400-3935 m.
Colombia (Norte de Santandér to Meta), Venezuela (Tachira). Map 3.
COLOMBIA. Boyaca: Paramo de La Rusia, NW. to N. of Duitama, Serrania
Pena Negra, Hoya de la Laguna Agua Clara, 3935 m, 10.xii.1972 (fl), Cleef 6992
(COL, U); Vado Hondo, Siberia, between Pefia de Arnical and Alto de Mégotas,
3290 m, 4.iv.1973 (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: Péaramo de Sumapaz, Hoya El Nevado, Laguna El Sorbedero,
3550 m, 1.ii.1972 (fl), Cleef 1489A (COL). Norte de Santandér: Paramo de Hatico
between Toledo and Pamplona, c. 2900 m, 12—13.i11.1927 (fl), Killip & Smith 20661
(K, NY, US). Santandér: Paramo del Almorzadero, 3600-3800 m, 28.xi.1941 (fl),
Cuatrecasas 13496 (F, NY, P); Paramo Rico, near Vetas, 3750-3850 m, 18.i.1927
(fl), Killip & Smith 17659 (NY).
THE GENUS HYPERICUM L.
Map3 4. H. garciae @;5.H. acostanum @.
VENEZUELA. Tachira: Paramo de Tama, El Paramito, 2550 m, viii.1939 (fl),
Chardon 81 (VEN).
H. garciae differs from the coriacous-revolute-leaved form of H. phellos ssp. phellos
(‘patens’) principally by its narrower, closely revolute leaves, a character that
distinguishes it from all other species in sect. Brathys, except H. acostanum and 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 affinis, sed foliis patulis persistentibus, pro ratione angustioribus,
uninervis, margine revolutis, sepalis angustioribus, petalis longioribus, inter alia
differt. Type: Ecuador, Loja, Cant6n 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,
zy
NORMAN K. B. ROBSON
scarcely tetrastichous, persistent, not articulated but sometimes breaking off above
base; lamina 8-22 x 0-5—2-5 mm, narrowly oblanceolate to linear, broadly to
narrowly revolute, sometimes concealing lower surface apart from midrib, not
carinate, paler or ferrugineous and smooth beneath, lucent above, not glaucous,
coriaceous; apex acute, base parallel-sided, not sheathing, pairs united to form a
narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense, impressed.
Inflorescence 1-flowered, terminal, often with pseudo-dichotomous branches;
pedicel 3-15 mm long, not upwards incrassate; upper leaves foliose. Flowers 20-25
mm in diam., stellate. Sepals 6-9 x 1-1-5 mm, narrowly triangular-lanceolate to
oblong-linear, acute, sometimes revolute above; veins 5—7, obscure, not visibly
branching, midrib slightly prominent; glands linear, distally punctiform. Petals
bright (?) yellow, tinged red in bud, 10-14 x 5—7 mm, c. 2 X sepals, obovate; glands
linear and striiform. Stamens c. 100, longest 4-5 mm long, c. 0-4 x petals. Ovary c. 2
x 1mm, 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 Roldés memorial monument, 2700 m, 19.11.1985 (fl & fr), Harling & 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. inJ.
Wash. Acad. Sci. 48: 314 (1958); Rodriguez Jiménez in Reitz, Fl. Ill. Catarin.,
Hipericaceas: 5, fig. 1 B (1980). Type: Uruguay, Mardonado, Pan de Aztcar cerca
del Castillo de Piridpolis, xi.189- (fl & fr), Arechavaleta (MVM, holotype; GH!,
US!, photographs).
H. tenuifolium St.-Hil. in St.-Hil., Jussieu & Cambess., Fl. Bras. Merid. 1: 337
(1828); Walpers, Repert. Spec. nov. 1: 390 (1842); Dietrich, Syn. PI.: 1236 (1847);
Reichardt in Martius, Fl. Brasil. 12: 188 (1878), non Pursh (1814). Type: Brazil,
Sao Paulo, prope urbes Sancti Pauli, Villa do Castro et Curityba, iii. 1820 (fl & fr),
St.-Hilaire 1174 (P!, holotype; US!, photograph).
H. hilaireanum L.B. Sm. in J. Wash. Acad. Sci. 48: 314 (1958); Angely, Fl. Anal.
Paranda: 451 (1965). Type as for H. tenuifolium St.-Hil.
Subshrub or perennial herb 0-07-0-35 m tall, erect, with taproot, branches few, strict
or decumbent, lateral, from base or lower half of stem. Stems green, 4-lined above,
eventually terete, gland-dotted, with + prominent glands along raised lines; cortex
eventually exfoliating in strips; internodes 3-21 mm long. Leaves sessile, outcurving
and deflexing, not tetrastichous, persistent, not articulated but occasionally breaking
off above base; lamina 10-40 x 1—2 mm, linear, revolute, concealing all or most of
lower surface apart from midrib, not carinate, paler beneath?, dull above, +
glaucous, subcoriaceous; apex acute to acicular, base parallel, not sheathing, pairs
united to form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands
dense, slightly impressed or not. Inflorescence 1—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
xX 1-5-2-5 mm, linear-lanceolate to lanceolate or narrowly oblong, acute to acumin-
ate, margin slightly revolute or plane; veins 3-5, unbranched, midrib slightly
prominent or not; glands linear towards base or wholly punctiform. Petals bright (?)
yellow, not (?) tinged red in bud, 7-12 x 3-4-5 mm, 0-8-1-2 x sepals, obovate-
oblong; glands linear, distally punctiform. Stamens 25—60, longest S—8(—10) mm, c.
0-75 x petals, 3(5)-fascicled, the double (antesepalous) fascicles sometimes incom-
THE GENUS HYPERICUM L.
pletely united, with filaments within each fascicle united shortly or usually to 0-7 of
their length. Ovary 1-5—4 x 1-1-5 mm, narrowly ovoid to narrowly ovoid-ellipsoid;
styles 3, 2-4 mm long, c. 1-1-3 X ovary, outcurving; stigmas subcapitate. Capsule
5-7 X 2-2-3 mm, ovoid-cylindric to ellipsoid, shorter than sepals. Seeds 0-8-1 mm
long, ecarinate; testa finely scalariform.
In dry, stony or damp grassland; lowland to 1000 m.
Brazil (SAo Paulo, Parana, Santa Catarina, Rio Grande do Sul), Uruguay. Map 4.
BRAZIL. Sao Paulo: Sao Paulo to Villa Emma, xii. 1933 (fl), Brade 12978 (MO).
Parana: Municipio Guarapuava, estrada para Laranjeiras do Sul, 15.xi.1957 (fl),
Hatschbach 4253 (US); Municipio Ponta Grossa, Parque Vila Velha, Furnas,
i.x.1965 (fl & fr), Hatschbach 12854 (K). Santa Catarina: Municipio Abelardo Luz,
8-12 km N. of Abelardo Luz, 900-1000 m, 15.xi.1964, Smith & Klein 13305 (NY,
US); Municipio Xanxere, 11 km N. of Abelardo Luz, 500-600 m, 25.xii.1956, Smith
& Klein 9231 (US). Rio Grande do Sul: Vila Manresa, prope Porto Alegre,
21.x.1951, Rambo 51325 (US); Porto Alegre, Morro da Policias, 21.ii.1902 (fl & fr),
Malme Regn. II 1408 (S).
URUGUAY. Mardonado: see type. Lavalleja: Co. Penitente, 10.1.1909 (fl),
Berro 5533 (US).
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.
Ze
30
NORMAN K. B. ROBSON
I agree with Rodriguez Jiménez (1980) that H. piriai cannot be divided into a
northern species (H. hilaireanum) and a southern one (H. piriai sensu stricto). The
more northern ones, suffruticose with fewer larger flowers and longer sepals, are the
nearest morphologically to H. acostanum.
7. Hypericum irazuense Kuntze [Rev. Gen. Pl: 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. Boissier 6: 258 (1898), in op. cit. 11, 8: 177 (1908), in Engler & Prantl,
op. cit., 2nd ed. 21: 181 (1925), non Juss. (1804).
Icon: Fig. 5A.
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 Xx 1-5-3-5 mm, narrowly elliptic to oblan-
ceolate, plane to recurved, not carinate, concolorous, not glaucous, subcoriaceous to
coriaceous; apex acute, base cuneate, not sheathing, pair free but with incomplete
narrow interfoliar ridge; basal veins 3-5, parallel, inner pair (of 5) branching above,
tertiary reticulation not visible; laminar glands rather dense, impressed above, not
prominent beneath. Inflorescence 1-flowered, terminal and on short lateral shoots,
with extension shoots from 2—3 nodes below; pedicel 2-5—4 mm long; upper leaves
bracteose. Flowers 25-30 mm in diam., stellate. Sepals 7-5-9 x 1-5-2 mm, linear-
lanceolate, acute; veins 5, unbranched, midrib not prominent; glands mostly linear,
punctiform near apex. Petals bright yellow to orange-yellow, not (?) tinged red,
13-18 x 7-10 mm, c. 2 X sepals, oblanceolate to obovate; apiculus acute; glands
linear, interrupted distally. Stamens c. 100, longest 6-9 mm long, 0-35-0-5 x petals.
Ovary 1-5—2-5 x 1-1-5 mm, ovoid; styles 3, 9-10 long, 4-6 x ovary, free, spreading,
distally incurved; stigmas small. Capsule 6-7 x 4-5 mm, broadly ovoid or ellipsoid
to globose, shorter than sepals. Seeds 1-1-3 mm long, scarcely carinate; testa finely
scalariform. 2n = 12.
On open paramo slopes or among bamboo (Chusquea); 2700-3730 m.
Costa Rica (San José, Cartago, Lim6n), on the cordillera and on Volcan Irazi and V.
Turrialba, and in adjacent Panama (Chiriqui). Map 5 (p. 32).
COSTA RICA. Cartago: Volcan Irazt, 3000-3300 m, 5.v.1928 (fl), Stork 2011 (F,
MICH, NY); Finca Quemado to top of Volcan Turrialba, 2800-3200 m, 10.v.1971
(fl), Wilbur 14319 (DUKE); Cordillera de Talamanca, Cerro de la Muerte, Pan-
American Hwy., 5 km above Millsville, 3400-3500 m, 22.vii.1949 (fl), Holm & Iltis
458 (F, K, NY, P). Limon: Chirrip6 National Park, between Casa de Administracié6n
and peak, c. 3400 m, 13.11.1983 (fl & fr), Garwood et al. 1161 (BM). San José: 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.1968 (fl), Wilbur & Stone 10047 (DUKE, F, MO).
PANAMA. Chiriqui: Cerro Fabrega and vicinity near Costa Rican frontier,
3150-3335 m, 7—8.iv.1976 (fl), Weston 10189 (MO).
H. irazuense has apparently been derived from H. phellos subsp. oroqueanum in a
similar way to the derivation of H. stenopetalum from H. phellos subsp. phellos, but it
can be distinguished from H. stenopetalum both morphologically and geographi-
cally.
THE GENUS HYPERICUM L. 31
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SING win Sh
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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 12;b x 1; d—g, i-k x 3;c,h x 4). A. Khan, Vickery & Tebbs 1398; B. Luteyn 6032.
The record of H. 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
Map5 7. H. irazuense A; 8. H. stenopetalum @.
Venezuela, Mérida, 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 Annis Sci. nat. (Bot.) IV, 18: 297 (1862), nomen.
H. meridense Steyerm. in Fieldiana Bot. 28 (2): 394. (1952). Type: Venezuela,
Mérida, between San José 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—S, diverging, all or only midrib branching, tertiary reticula-
tion obscure or not visible; lamina glands rather dense, impressed above, +
prominent beneath. Jnflorescence 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 S—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, (S—)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 (Santandér), Venezuela (Tachira, Mérida). Map 5.
COLOMBIA. Santandér: [Péramo 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. 188169 (COL, US).
VENEZUELA. Mérida: Paéramo de Los Leones (La Lagunita, La Canada
Grande) al W. de Murucuba, 3400-3500 m, 31.v.1930 (fl), Gehringer 159 (F, NY,
VEN); Péramo de San José, monte El Cupio, 3000 m, 1.i1.1973 (fl & fr), Cuatrecasas,
Ruiz-Teran & L6épez-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.
9. Hypericum carinosum R. Keller
in Bull. Herb. Boissier, I, 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 bot. Club 56: 102—103 (1929). Type: Venezuela, Mérida,
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 Annis Sci. nat. (Bot.) IV, 18: 297 (1862)
pro parte, quoad syn. Trevirani.
H. stenoclados Cuatrec. in Ciencia Mex. 4: 64 (1943). Type: Colombia. Santandér,
Paéramo de Almorzadero, extremo sur, Peralonso, 3200 m, 19.vii.1940 (fi),
Cuatrecasas & Garcia Barriga 9919 (COL, holotype; F!, NY!, US! isotypes).
Shrub 0-6—c.2-5 m tall, erect, with branches strict, lateral, short (flowering) and long
(extension). Stems purplish-brown, 4-lined and ancipitous when young, eventually
terete, without epidermal emergences, cortex exfoliating in strips; internodes 2—9
mm long. Leaves sessile, spreading from the base or ascending but not imbricate or
tetrastichous, persistent; lamina 3-5—5 x 0-8-2 mm, narrowly elliptic or narrowly
oblong to oblanceolate-spathulate, plane, rarely apically subconcave, markedly
carinate, concolorous, not or slightly glaucous, chartaceous to subcoriaceous; apex
acute to subacute, base cuneate to angustate, not sheathing, pairs almost free
(interfoliar ridge rudimentary); basal vein 1, unbranched; laminar glands dense, not
impressed above, prominent beneath. Jnfloresence 1—-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 Santandér, Santandér), Venezuela (Mérida). Map 6 (p. 37).
COLOMBIA. Norte de Santandér: environs of Pamplona. 10.x.1943 (fl), de
Garganta 626 (F). Santandér: vicinity of Las Vegas, 2600-3600 m, 21—23.xii.1926,
Killip & Smith 15921 (GH, NY, US); vicinity of California, 2800 m, 11—27.i.1927,
Killip & Smith 16904 (F, GH, NY, US).
VENEZUELA. Mérida: Paramo de Los Leones, 3400-4500 m, 31.v.1930 (st),
Gehringer 149 (VEN).
H. carinosum is related to H. phellos subsp. phellos ‘diversicaule’, and its area of
distribution is to the south-east of that of the latter. The two taxa, however, are quite
3D
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 H. carinosum, but these differences do not merit taxonomic
recognition.
10. Hypericum simonsii N. Robson, sp. nov.
H. phellos affinis, sed caulis haud suberosis, foliis persistentibus deflexis oblongis
apice rotundatis basi conjunctis venatione valde reticulata, sepalis subacutis vel
rotundatis, stylis (S—)6—9 mm longis, differt. Type: Colombia, Magdalena, Sierra
Nevada de Santa Marta, 1880 (fl), Simons s.n. (BM!, holotype).
H. stenopetalum [var.] B 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.i1.1959 (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 S—7 basal or near-basal veins.
11. Hypericum papillosum N. Robson, sp. nov.
H. simonsii N. Robson affinis, sed habitu humiliore densiore, foliis sepalisque
subtiliter papillosis, foliis angustioribus apice obtusis vel rotundatis basi breviter
vaginatis, stigmatibus valde capitatis, differt. Type: Colombia, Boyaca, Sierra
THE GENUS HYPERICUM L. 35
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S pS a aaa
px
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Ne
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 12; 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 2728A.
Nevada del Cocuy, above Laguna Grande, c. 3900 m, 14. viii.1957 (fl), Grubb, Curry
& Fernandez-Perez 401 (K!, holotype; US!, isotype).
Icon: Fig. 6B.
Shrub to c. 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,
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/Paéramo de Chita. Map 2 (p. 24).
COLOMBIA. Boyaca: Paramo de Pisva, road between Socha and La Punta, Km
61, 5-5 km E. of Los Pinos, 3680 m, 14.vi.1972 (fl), Cleef 4541 (U); same road, Km
70, 1-5 km, W. of El Cadillal, 3530 m, 16.vi.1972 (fl), Cleef 4631 (U); Paramo de
Chita, vertiente oriental, sitio Piedras Negras, 3500-3700 m, 13.vii.1967 (fl), Jara-
millo & van der Hammen 2728A (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 [Zipaquir4], viii.1855, (fl), Triana 5464 (COL?, holotype;
BM!).
H. gyropodioides sensu Jackson & Hooker, Index Kew. 1: 1193 (1893), sphalm.
Shrub 0-8—2:-5 m tall, erect, with branches strict to ascending, lateral, short (flower-
ing) and long, narrowly spiciform to cylindric (extension). Stems dark red-brown,
4-lined when young, eventually terete, without epidermal emergences, cortex
exfoliating in strips or small flakes; internodes 1-5—2-5 mm long. Leaves sessile,
imbricate to spreading, eventually reflexed, markedly terastichous; lamina 3-6 X
1-5-3 mm, oblong to elliptic or lanceolate, incurved-conduplicate, midrib usually
prominent beneath, margin narrowly hyaline and plane or undulate, not glaucous,
subcoriaceous; apex acute to acuminate, slightly cucullate, base broadly cuneate to
angustate, loosely clasping, pairs united to form narrow interfoliar ridge; basal vein
1, sometimes branched; laminar glands dense, not impressed or prominent, visible
on both sides. Inflorescence 1-flowered, terminal and on short lateral branches;
pedicels absent; upper leaves foliose, appressed to calyx. Flowers (15—)25—30 mm in
diam., stellate. Sepals (S—)6—-7 X (2—)2:5-3 mm, ovate-oblong to ovate-lanceolate,
acute to obtuse, + subcucullate, margin hyaline; veins 7, unbranched, midrib
scarcely prominent; glands linear, interrupted distally. Petals bright yellow, 13-16 x
9-10 mm, 2-2-2 X sepals, very obliquely obovate; apiculus subacute; glands
striiform and punctiform. Stamens c. 120, longest 6-8 mm long, c. 0-5 X petals.
Ovary 2-5-3 X 1-5-2 mm, ellipsoid; styles 3, 4-5 mm long, c. 1-4 X ovary,
spreading-incurved; stigmas small to narrowly capitate. Capsule c. 6 X 3-5 mm,
ellipsoid, shorter than sepals. Seeds 1-1-2 mm long, ecarinate; testa finely reticulate-
scalariform.
In subpaéramo and paramo, most frequently in thickets; 2850-4100 m.
THE GENUS HYPERICUM L.
Map6 9.H. carinosum A; 12. H. lycopodioides O; 13. H. thuyoides e.
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
José, 3450 m, 9.vi.1973 (fl), Cleef 10117 (U). Boyaca: Sierra Nevada del Cocuy,
Valle de San José, c. 4000 m, 10.ix.1957 (fl), Grubb, Curry & Fernandez-Perez 768
(COL, K, US); between Chita and Sacama, Quebrada del Curial (La Porquera),
3350 m, 15.ix.1969 (fl), Cuatrecasas & Laureano Rodriguez 27787 (BM, COL, US).
Cundinamarca: see type.
H. lycopodioides is morphologically and geographically intermediate between 10 H.
simonsii and 13 H. thuyoides. It is very similar to the latter, differing essentially in the
stiff, acute to acuminate leaves that give the shoots an Araucaria-like appearance.
13. Hypericum thuyoides Kunth
in Humboldt, Bonpland & Kunth, Nova Gen. et Sp. Pl. 5: 187, t. 456 (1822); R.
Keller in Bull. Herb. Boissier Il, 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 Ibagué, x.1805
(fl), Humboldt & Bonpland (P-HUM!, holotype; B-WILLD, P!, isotypes).
Brathys thuyoides (Kunth) Spach, Hist. nat. vég. Phan. 5: 448 (1836), in Annls Sci.
nat. (Bot.) II, 5: 366 (1836).
H. hartwegii var. patens R. Keller in Bull. Herb. Boissier Il, 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. et Sp. Pl. 5: t. 456 (1822).
Shrub or small tree (0-5—)1—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 xX 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
37
38
AN
NORMAN K. B. ROBSON
and usually undulate, not glaucous, thinly chartaceous; apex acute to obtuse,
cucullate, base cuneate to angustate or shortly pseudopetiolate, pairs united to form
marked interfoliar ridge; basal vein 1, branched or usually unbranched; laminar
glands + dense, visible on both sides or above only. Inflorescence 1-flowered,
terminal and on short lateral branches, the whole shoot cylindric to conic and
sometimes racemiform due to retarded maturity of terminal flower, rarely with
pseudo-dichotomous branches from node below; pedicel amost absent; upper leaves
foliose. Flowers 20-30 mm in diam., stellate. Sepals 3-5 x 1-5-2-5 mm, oblong-
lanceolate or elliptic to obovate-elliptic, acute to obtuse, margin narrowly hyaline;
veins 9-11, not or scarcely branched, midrib apically incrassate; glands linear,
punctiform distally. Petals bright yellow to orange-yellow, 10-17 x 6-11 mm, 3-4 x
sepals, very obliquely obovate; apiculus obtuse; glands striiform with few distal dots.
Stamens 100-120, longest 5-6 mm long, c. 0-4—0-5 x petals. Ovary 2-3 X 1-2 mm,
ovoid to ellipsoid; styles 3, 4-7 mm long, c. 2 X ovary, spreading; stigmas scarcely to
narrowly capitate. Capsule c. 3-5 X 2 mm, ellipsoid, shorter than sepals. Seeds not
seen.
In scrub and thickets on slopes of the paramo; especially in boggy areas; 2500-—
3600 m.
Colombia (Boyacaé, 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.1974 (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 985 (COL, U).
Tolima?: see type.
H. thuyoides is a rather variable species with a restricted distribution in central
Colombia, records from Narifo 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 (‘H. 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. 1A (1936). Type:
Colombia, Cundinamarca, Guasca, Los Gaques wood, 3000 m, 24.iv.1932 (fi),
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, Mérida, Paramo de Timotes (Jahn 847) (Reprium Spec.
nov. Regni veg. Beih. 43: 484 (1927)) must be a misidentification, but I have not seen the
specimen.
THE GENUS HYPERICUM L.
along shoot, oblong-ovate or ovate to triangular-ovate or subcircular, incurved-
conduplicate or saccate, with midrib slightly prominent beneath or not, margin
narrowly hyaline, not glaucous, chartaceous, deciduous almost at the base after
turning brown, leaving narrow basal ring; apex subacute or apiculate-obtuse to
rounded, base cordate-amplexicaul, pairs + perfoliately united; basal vein 1,
sometimes with 1-2 pairs of near-basal branches, rarely with visible tertiary reticu-
lum; laminar glands dense to rather few, large, + prominent on both sides.
Inflorescence 1-flowered, terminal and on short lateral branches, the whole shoot
cylindric to conical, without pseudo-dichotomous branches; pedicel 1-3-5 mm long;
upper leaves foliose. Flowers 20—40 mm in diam., stellate. Sepals 5-8 x 2-3-5 mm,
the outer oblong to ovate, often cordiform, the inner oblong to lanceolate, acute or
obtuse to rounded, cucullate; margin narrowly hyaline; veins 7-11, branching and
anastomosing distally, midrib distally incrassate; glands linear, distally punctiform.
Petals bright yellow, 10—20 x 6-12 mm, 2-2-5 X sepals, obovate; apiculus obtuse;
glands striiform and punctiform. Stamens c. 120-150, longest 5-6 mm long, c.
0-4—0-5 x petals. Ovary 2-5-3 X 1-5 mm, narrowly ovoid; styles 3, 5—8 mm long, c.
2-2-5 X ovary, spreading-incurved; stigmas narrowly capitate. Capsule 5-6 X 4mm,
broadly ellipsoid to subglobose, equalling or exceeding sepals. Seeds c. 1 mm long,
ecarinate; testa finely scalariform-reticulate.
In degraded forest and scrub on open slopes and banks in the paramo and subpara-
mo, mainly near water and sometimes in pure stands (Lozano-C. & Schnetter
(1976)); 2730-3590 m.
Colombia (Cundinamarca, Meta). Map 7.
Map7 14. H. goyanesii A; 16. H. cuatrecasii @; 17. H. quitense @.
Bp
40
NORMAN K. B. ROBSON
COLOMBIA. Cundinamarca: Péramo de Guasca, 3300 m, 16.xii.1938 (fl), Balls
5764 (BM, COL, K); Paramo de Zipaquira [Guerrero], between Zipaquiré and
Pacho, 3100-3200 m, 16.vi.1940 (fl), Cuatrecasas 9526 (COL, F, P); Paramo de Cruz
Verde, fondo del valle al sur de La Viga, 3590 m, 28. viii. 1972 (fl), Cleef 3287 (BM,
COL, U). Meta: Macizo de Sumapaz, alredores de la Laguna La Guitarra, 3380
—3420 m, 4.vii.1981 (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.1868? (fl), Stiibel
Colomb. 124 (Bt, holotype; F!, NY!, photographs).
H. myricariifolium Hieron. ex R. Keller in Bot. Jahrb. 42: 130 (1908), in Bull. Herb.
Boissier II, 8: 178 (1908). Type: as for H. myricariifolium Hieron.
H. lindenii sensu Cuatrec. in sched., non R. Keller.
Shrub up to 2 m tall, bushy or with branches ascending to spreading or prostrate and
matted, lateral, short (flowering) or long and narrowly conical (extension), rarely
pseudo-dichotomous. Stems orange-brown, 4-lined when young, sometimes wrink-
led or with corky emergences especially near flowers, eventually terete, cortex
exfoliating in irregular flakes; internodes 1-4—6 mm long. Leaves sessile, imbricate to
spreading, tetrastichous, cupressoid; lamina 1-3-5 x 0-7—2(—2-6) mm, triangular-
ovate to oblong-ovate, incurved-cucullate, with midrib plane or usually impressed
beneath, margin relatively broadly hyaline, not glaucous, coriaceous; apex subacute
to obtuse, base rounded to cordate-amplexicaul, pairs shortly united; basal vein 1,
unbranched, without visible tertiary reticulum; laminar glands dense to few, large,
prominent beneath or not. Inflorescence 1-floweted, not terminal, on short lateral
branches, the whole shoot narrowly conical, racemiform, rarely with a pseudo-
dichotomous branch; pedicel 1-1-5 mm long; upper leaves foliose. Flowers 18-25
mm in diam., stellate. Sepals 4-5 x 1-7-2-7 mm, the outer ovate, sometimes
cordiform, the inner ovate-lanceolate to lanceolate-oblong, acute to rounded,
cucullate, margin narrowly to rather broadly hyaline; veins 9-11, usually unbran-
ched, midrib not prominent beneath; glands linear to punctiform. Petals bright (?)
yellow, 10-15 x 6-9 mm, c. 2:2—3 X sepals, obovate; apiculus rounded; glands
striiform and punctiform. Stamens 60-80, longest 5—6 mm long, c. 0-35—0-5 x petals.
Ovary 2-3 X 1-1-5 mm, narrowly ovoid-ellipsoid; styles 3, 5—7 mm long, c. 2-2-5 x
ovary, spreading-outcurved; stigmas narrowly capitate. Capsules 5 x 4mm, 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 Usmé, Paramo de Chisa-
ca, c. 4120 m, 9-11.xi.1958 (fl), Barclay & Juajibioy 6178 (COL, MO, NY); Macizo
de Sumapaz, Andabobos, 3760-3720 m, 8.1.1969 (fl), Cuatrecasas & Jaramillo 27031
(BM, US); Paramo de Cruz Verde, W. slopes, 3150 m, 7.x.1938 (fl), Cuatrecasas 409
(COL, FE, 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 H. myricariifolium.
It is clearly a high-altitude derivative of H. goyanesii, and the altitudinal ranges of the
two taxa overlap from 3000—c. 3600 m; there is no evidence of hybridization between
them. They can be distinguished by the leaf shape (see key), which gives the young
THE GENUS HYPERICUM L.
shoots of H. myricariifolium a cupressoid appearance lacking in H. goyanesii, the
leaf aspect (leaves becoming reflexed in H. goyanesii but no more than spreading in
H. myricariifolium), and usually by the leaf size. It seems advisable, therefore, to
recognize them as species, despite their overlapping ranges.
16. Hypericum cuatrecasii Gleason
in Revta Acad. Colomb. 5: 38 (1942), in Bull. Torrey bot. Club 70: 174 (1943). Type:
Colombia, Boyaca, Cordillera Oriental, between Arcabuco and Tunja, 2950 m,
5.viii.1940 (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, 9-125 X 6-5—7-5 mm, c. 1-8 X sepals, obliquely obovate; apiculus obtuse;
glands mostly striiform, distally punctiform. Stamens c. 120-150, longest 5-6 mm
long, c. 0-5 X petals. Ovary 2-5-3 x 1-5-2 mm, broadly ovoid; styles 3(4), 5-6-5 mm
long, 2 X ovary, spreading-incurved; stigmas scarcely to narrowly capitate. Capsule
c.6 X 3-5 mm, ellipsoid, slightly exceeding sepals. Seeds not seen.
In paramo and degraded forests; 1990-3000 m.
Colombia (Boyaca, Santandér), in the Paramo de Arcabuco. Map 7 (p. 39).
COLOMBIA. Boyaca: Sachica, 1990 m, viii.1964, Saravia 4425 (COL). Santan-
dér: 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 Santandér.
17. Hypericum quitense R. Keller
in Bull. Herb. Boissier I, 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 Biblian, c. 3400 m, 13.viii.1933, Diels 497 (B+, holotype).
Icon: Fig. 7.
Shrub or shrublet 0-3--1(-1-5) m tall, erect or spreading, with branches divergent,
lateral, short (flowering) and long (extension), not pseudo-dichotomous. Stems
reddish-brown, 4-lined and markedly ancipitous when young, eventually terete,
without wrinkles or corky emergences, cortex exfoliating in flakes; internodes 4—10
41
42 NORMAN K. B. ROBSON
oH “a
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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 4; b, f, g x 4; c—-e x 6). All Camp 4133.
mm long. Leaves sessile, spreading to deflexed, obscurely tetrastichous, turning
brown, deciduous with the cortex; lamina 4—9-5 x 2-5—5 mm, sometimes increasing
in size along shoot, oblong (lower) to broadly ovate, plane or slightly saccate or
conduplicate, midrib prominent beneath, margin narrowly hyaline, otherwise pale
green but not or only slightly glaucous, subcoriaceous; apex acute (lower) to obtuse,
base broadly cuneate, pairs shortly united; basal or near-basal veins 3(5), with
laterals and sometimes midrib branching, tertiary reticulum obscure or apparently
absent; laminar glands mainly peripheral, dense, rather large, not prominent.
Inflorescence 1-flowered, terminal and on very short lateral shoots, the whole shoot
narrowly cylindrical, without pseudo-dichotomous branches; pedicel 3—7 mm long;
upper leaves foliose. Flowers 8-12 mm in diam., stellate. Sepals 5 x 1-2 mm, elliptic
to narrowly oblong, the outer sometimes broader, apiculate-obtuse to acute, sub-
cucullate, margin narrowly hyaline; veins 3-5, not or scarcely branching, all promin-
ent; glands all linear or distally punctiform. Petals bright yellow, 5-6-5 x 3-4 mm,
1-5-2 x sepals, obovate; apiculus subacute to obtuse; glands few, striiform and
THE GENUS HYPERICUM L.
punctiform. Stamens 30-45, longest 3—4 mm long, c. 0-65 x petals. Ovary 1-5-2 x
0-7-1 mm, narrowly ovoid; styles 3, 3-4 mm long, c. 2 X ovary, spreading; stigmas
broadly capitate. Capsule 3-4-5 x 2-5—3-5, ellipsoid, equalling or slightly exceeding
sepals. Seeds c. 0-7 mm long, ecarinate, testa finely scalariform.
In damp meadows of the paramo; 2600-3800 m.
Ecuador (Bolivar, Tungurahua, Cafar, Azuay). Map 7 (p. 39).
ECUADOR. Tungurahua: around Lago Pisayambo, 3800 m, 26.xii.1964 (fl),
Knight 268 (BM). Bolivar: Cordillera Occidental, Pucara de Telimbela, 2600-3000
m, 18.xi.1943 (fl), Acosta Solis 6825 (F). Canar: near Pimo, 3060-3120 m, 9.vii.1945
(fl), Camp E-4133 (BM, NY). Azuay: mountains above Sayausid, 3000-3200 m,
18. iii. 1974 (fl & fr), Harling & Anderson 12630 (GB).
H. quitense is related to H. cuatrecasii but is much smaller in all parts.
18. Hypericum loxense Bentham
Pl. Hartweg.: 126 (1843); R. Keller in Bull. Herb. Boissier II, 8: 177 (1908), in Engler
& Prantl, Nat. Pfanzenfam. 2nd ed. 21: 181 (1925); Gleason in Bull. Torrey bot.
Club 56: 102 (1929). Type: Ecuador, Loja, ‘in montibus Loxa’, viii.1842? (fl & fr),
Hartweg 720 (K!, holotype; BM!, F!, P!, isotypes).
Shrub or shrublet, 0-2—1(—1-5) m tall, erect or decumbent to prostrate, with branches
ascending to strict, lateral, short (flowering) and long (extension), not pseudo-
dichotomous. Stems reddish-brown, 4-lined and ancipitous when young, eventually
terete, without wrinkles or emergences, cortex exfoliating in irregular flakes or
strips; internodes 1-5—10 mm long. Leaves sessile, erect, imbricate and sometimes
markedly tetrastichous when young, spreading and turning brown, deciduous with
the cortex (or sometimes breaking off earlier); lamina 2-10 x 0-6-3 mm, not
increasing in size along shoot, narrowly oblong or oblanceolate or linear to triangu-
lar-lanceolate or triangular-ovate, incurved-conduplicate or subsaccate, midrib
prominent beneath or not, margin narrowly hyaline, not or slightly glaucous,
subcoriaceous to chartaceous; apex acute to obtuse, base narrowly cuneate or
parallel-sided to broadly cuneate or truncate, pairs shortly united; basal veins
1(3-5), with midrib sometimes branching, tertiary reticulum rarely present; laminar
glands dense to rather sparse, varying in size, prominent or not. Inflorescence either
(i) wholly 1-flowered, terminal and on very short lateral shoots, the whole shoot
narrowly cylindric, with lateral branches sometimes also bearing flowering branches
but rarely with pseudo-dichotomous branches, or (ii) 1—c. 15-flowered in terminal
and sometimes lateral dichasia or mixed dichasia and pseudo-dichotomies; peduncle
and pedicels 2—9 mm long; upper leaves foliose, bracts gradually smaller. Flowers
6-10 mm in diam., stellate. Sepals 1-5—5-3 x 0-5—1-6 mm, elliptic or lanceolate to
narrowly oblong, the outer sometimes broader, apiculate-obtuse to acute, some-
times cucullate, margin narrowly hyaline; veins 3—S, not or scarcely branching, not
or only the midrib slightly prominent; glands all linear or distally punctiform. Petals
pale or bright yellow to orange, darker or reddish in bud, 3—6(—7-5) x 1-5-3 mm,
1-5-2 X sepals, obovate; apiculus obtuse or obsolete; glands few, striiform and
punctiform. Stamens 20—40, longest 2-4 mm long, c. 0-65 X petals. Ovary 0-7—1-5 x
0-5-1 mm, narrowly ovoid; styles 3(4), 1-3-3 mm long, 1-3-2 x ovary, spreading;
stigmas + broadly capitate. Capsule 2-5-4 x 1-5-2-2 mm, ellipsoid or ovoid-
ellipsoid, equalling sepals. Seeds 0-7—0-8 mm long, ecarinate, testa finely scalar-
iform.
On dry slopes and in rocky places in the paramo; 2350-3800 m.
Ecuador (Cotopaxi, Chimborazo, Canar, Azuay, Loja), Peru (Cajamarca, La
Libertad). Map 8.
H. loxense is closely related to H. quitense, which although overlapping it in
distribution is apparently ecologically distinct. In addition, H. loxense has narrower
and/or smaller leaves and usually smaller flowers.
H. loxense comprises two populations of which subsp. aequatoriale is nearer to H.
ne
TEeeea es
FALE LASY
ii
a
43
=
sed)
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44
NORMAN K. B. ROBSON
[ 80
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, Cafar, 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. Cafiar, between Azogues and Péaramo
de Huairacaja, 2700-3000 m, ix—x.1905? (fl & fr), Lehmann 4795 (B+, 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.
Ecuador (Cotopaxi, Chimborazo, Canar, Azuay, Loja).
ECUADOR. Cotopaxi: Coraz6n, 3200 m, 22.vi.1876, André K. 436 (K). Chim-
borazo: Péaramo de Cachenco, c. 3800 m, 27.vii.1959 (fl), Barclay & Juajibioy 8258
(NY). Cafiar: near Pimo, 3060-3120 m, 9.vii.1945 (fl), Camp E-4136 (BM, NY);
Tipococha, 3400 m, 9.vii.1939 (fl & fr), Penland 994 (F). Azuay: Benos, SW. of
Cuenca, c. 2700 m, 12.v.1974 (fl), Harling & Anderson 14561 (GB); San Miguel to
Cuenca, 25 km, 3000 m, 15.ix.1969, B. & C. Maguire 61712 (BM, NY). Loja:
between Loja and Cuenca, c. 20 km N. of Saraguro, 3080 m, 21.x.1980 (fl & fr),
Croat 50854 (BM, MO).
18b. Hypericum loxense subsp. loxense
Leaves spreading or more rarely erect and subimbricate; lamina 4-10 x 0-6-3 mm,
narrowly oblong or narrowly elliptic or rarely oblanceolate to linear, plane to
incurved, chartaceous, apex acute to rarely subacute, base narrowly cuneate to
parallel-sided, midrib often prominent beneath. Inflorescence branches usually
dichasial or mixed, more rarely 1-flowered. Sepals 2—4-5(5-3) x 0-5—1-2(1-6) mm,
narrowly elliptic to oblong, acute.
Ecuador (Chimborazo, Cafiar, southern Azuay, Loja), Peru (Cajamarca, La Liber-
tad).
ECUADOR. Azuay: Paramo Portete, 3600 m, 13.iii.1953 (fl), Prescott 793 (NY).
Chimborazo: Urbina, towards Mt. Chimborazo, c. 3700 m, 27.vii.1939 (fl & fr),
Asplund 7913 (S). Cafiar: near E] Tambo (c. 69 km S. of Sibombé), 2850-3000 m,
5.vii.1945 (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 (fi
& fr), Jameson 127 (K, W).
PERU. Cajamarca: above Sunchubamba, 3680 m, 8.vi.1957 (fl), Ellenberg 1862
(U). La Libertad: Playapampa, c. 2700 m, 15—24.vi.1923 (fl), Macbride 4525 (NY).
The disjunct populations in Chimborazo and northern Azuay differ in their decum-
bent to prostrate habit, whereas the collection from Cajamarca differs in having
oblanceolate subacute leaves and somewhat larger flowers than normal.
19. Hypericum sabiniforme Trev.
Hyper. Animad.: 15 (1861); Triana & Planchon in Annis Sci. nat. (Bot.) IV, 18: 296
(1862) [‘sabinaeforme’|. Type: Colombia, Boyaca, Prov. de Tunja, Soata, 1300 m,
iv.1843 (fl), Linden 1328 (BHU?, holotype; BM!, GH!, K!, P!, W!, isotypes).
Hypericum thymifolium sensu Turcz. in Bull. Soc. Nat. Moscou 31 (1): 386 (1858),
non Banks & Solander (1794) nec Kunth (1822).
Hypericum hartwegii sensu Triana & Planchon, tom. 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 I, 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!, P!, isotypes).
Hypericum trianae N. Robson in Cleef, Veg. Param. Colomb. Cord. Or.: 164, 305
(1981), nomen.
Shrub 0-3—0-5 m tall, erect, bushy, with branches erect, + strict, lateral or rarely
pseudo-dichotomous. Stems orange-brown, 4-lined when young, eventually terete,
cortex exfoliating in strips; internodes 1-5-4 mm long. Leaves sessile or with
pseudopetiole up to 1 mm long, outcurving to appressed, loosely to densely
imbricate, sometimes markedly tetrastichous, deciduous at or sometimes slightly
above the base; lamina 1-5-7 Xx 0-4-3 mm, ovate or elliptic to oblanceolate or
narrowly oblong, + cucullate, incurved, with margin narrowly hyaline, midrib
impressed beneath, concolorous, glaucous, coriaceous, markedly papillose or sub-
papillose above, less so beneath; apex obtuse to rounded, base cuneate-
45
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 xX 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 Susacén, paramo de Guantiva, 3140 m,
8.viii.1958 (fl), Jaramillo & Hernandez 965 (US); NW. of Belen, Quebrada Minas,
Hoya Clla. 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. 1973 (fl). Cleef, Cuatrecasas & Jaramillo
9236B (COL); Paramo de Gina, 3200 m, 17.vii.1940 (fl), Cuatrecasas & Garcia
Barriga. 9790 (F); Andes de Bogota, 2500-3000 m, (fl), Triana s.n. (BM).
The type of H. sabiniforme, which superficially resembles a species of the African
ericaceous genera Philippia or Blaeria, represents the extreme form of an eastward
a9
|
Map9 19. H. sabiniforme A; 22. H. hartwegii O; 23. H. maguirei @; 25. H. gleasonii @.
THE GENUS HYPERICUM L.
cline. At the western end of the cline is a form with much larger ovate to elliptic
pseudopetiolate leaves, markedly papillose above, and much larger flowers. Varia-
tion along the cline, however, appears to be continuous. The western form is near the
broad-leaved form of H. thuyoides, but differs in having thicker, glaucous leaves with
undulate to papillose epidermis.
Wood 3682 (Cundinamarca, between Neusa and San Cayetano) is intermediate
between 13 H. thuyoides ‘genistoides’ (see p. 38) and 19 H. sabiniforme.
20. Hypericum laricifolium Juss.
in Annls Mus. Hist. nat. Paris 3: 160, t.16 f.1 (1804); Choisy, Prodr. monogr. fam.
Hypéric.: 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), J. de Jussieu*
in Herb. A. L. de Jussieu 11800 (P — JUSS!, holotype; C!, F!, P!, isotypes).
H. acerosum Kunth in Humboldt, Bonpland & Kunth, Nova Gen. et Sp. Pl. 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. vég. Phan. 5: 366 (1836).
B. acerosa (Kunth) Spach, Hist. nat. vég. Phan. 5: 449. 1836, in Annls Sci. nat. (Bot.)
II, 5: 366 (1836).
Hypericum laricifolium var. acerosum (Kunth) Weddell, Chloris Andina 2: 271
(1857).
H. platypetalum Turcz. in Bull. Soc. Nat. Moscou. 31 (i): 388 (1858), non (Webb)
Steudel (1840). Type: Ecuador, Pichincha, Quito, 1836? (fl), Jameson 60 (LE,
holotype; BM!, K!, W!, isotypes).
H. racemulosum Turcz. in Bull. Soc. Nat. Moscou. 36 (1): 580 (1863). Type: Peru.
Amazonas, Chachapoyas, 1835 (fl), Mathews 1611 (LE, holotype; BM!, KW!, P!,
isotypes).
H. laricoides Gleason in Bull. Torrey bot. Club 56: 104. (1929). Type: Venezuela,
Mérida, Paéramo de Timotes, Péramos de Chachapos, 3800—4200 m, 24.i.1928 (fi),
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 (0-1—)0-3—3(—6) m tall, bushy or lax and spreading, with branches
erect to spreading, lateral, short (flowering) and long, narrowly to broadly conical
(extension). Stems 4-lined when young, eventually terete, blackish-brown, without
epidermal emergences, cortex exfoliating in strips or irregular flakes; internodes
1-5-5 mm long. Leaves sessile, spreading or erect, sometimes imbricate, not
markedly tetrastichous, persistent; lamina (2—)3—6(-8) x 0-3-2-2 mm, narrowly
elliptic or narrowly elliptic-oblong to acerose, conduplicate to incurved, with midrib
impressed beneath, margin narrowly hyaline and plane or subundulate, not
glaucous, chartaceous to coriaceous; apex subacute to obtuse, cucullate, base
narrowly cuneate to angustate, loosely clasping or not, pairs united to form narrow
interfoliar ridge; basal vein 1, not or obscurely branched; laminar glands dense,
scattered or in 1 row, impressed, usually visible beneath only. Inflorescence 1-
flowered, on short lateral branches, rarely branching pseudo-dichotomously; pedicel
1-5-5 mm long; upper leaves foliose. Flowers 15—25(-—30) mm in diam., stellate.
Sepals 3-7 x 0-7—2-2 mm, oblong or elliptic to narrowly ovate or lanceolate,
subacute to acute, cucullate, margin hyaline; veins 7-9, not or scarcely branched,
midrib distally incrassate; glands linear, distally punctiform. Petals bright golden
yellow to orange-yellow, 7-14)-22) x 4-10 mm, 2-3 x sepals, very obliquely,
obovate; apiculus obtuse; glands striiform and punctiform. Stamens 70-110, longest
4—7 mm long, c. 0-5 X petals. Ovary 2-3-5 x 0-8-1-5 mm, ellipsoid; styles 3, 4-9 mm
long, 2—4-5 x ovary, spreading; stigmas scarcely to narrowly capitate. Capsule 4—7 x
* The only specimen in Herb. de Jussieu that could be type material is labelled ‘Perou, 1780,
Dombey.’
48 NORMAN K. B. ROBSON
YY
Yirr Soy
; CO Yi
WW VF
f
Fig.8 A. H. laricifolium: (a) habit; (b) stem with leaves (Ecuador); (c) stem with leaves (Venezuela); (d)
leaf; (e) sepal; (f) petals; (g) stamens (partly cut away) and ovary; (h) capsule. B. H. martense: (i) habit;
(j) stem with leaves; (k) leaf; (1) sepal; (m) petal; (n) capsule (a, i x ¥2; b,c, j X 2; d—h, k—-n x 4). A.
Luteyn 6751 except (c) Luteyn 7583; B. Seifriz 421.
3-4 mm, ovoid to subglobose, equalling or exceeding sepals. Seeds 0-8—1 mm long,
ecarinate; testa finely scalariform.
In the subpéramo and pdramo, in open or more usually sheltered habitats, well-
drained or not; 2200—4300 m.
From western Venezuela (Lara, Trujillo, Mérida, Zulia) along the Cordilleras
THE GENUS HYPERICUM L.
Central and Oriental of Colombia and Ecuador to central Peru (Hudnuco, Ancash).
Map 10.
COLOMBIA. Antioquia: between Medellin and Retiro, around Las Palmas, c.
2800 m, 16.x.1947 (st), Gutiérrez, Barkley & Rosano 24(F). Boyaca: Paramo La
Rusia, NNW. of Duitama, 3550 m, 11.ix.1969 (fl & fr), Cuatrecasas & Rodriguez
27730 (BM, COL, US). Caldas: Rio Otin, 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
Puracé, S. of the Volcan, San Francisco, 3450-3400 m, 23.vii.1943 (fl), Cuatrecasas
14571 (COL, F, P). César: Municipio de Valledupar, Corregimiento de Manaure,
2800 m, n.d. (fl), Rodriguez s.n. (COL). Chocé: Macigo de Tatama, valle El
Encanto, 3420 m, 13.11.1983 (fl), Cleef & Salamanca s.n. (COL). Cundinamarca:
Paramo de Guasca, vicinity of Bogota, 3150 m, 23.viii.1959 (fl), B. & C. Maguire
44029 (BM, NY); Paramo de Palacio, Lagunas de Buitrago, 3550 m, 27.iv.1973 (fl),
Cleef 9577 B (COL). Magdalena: Sierra de Perija, plain between Cerro Venado and
Cerro Avi6n, 3270-3350 m, 8.xi.1959 (fl), Cuatrecasas & Romero-Castaneda 25128
(COL). Narifo: Volcan de Cumbal, via Laguna Bolsa, 2740-2980 m, 11.ix.1944 (fl),
Ewan 16139 (BM, S). Norte de Santandér: Ocania to Pamplona, Serra P1. , 2400-3300
m, 30.viii.1878 (fl), Kalbreyer 711 (K), Putumayo: Lagunda de la Cocha, Quebrada
de Santa Lucia, 2850 m, 8.1.1941 (fl), Cuatrecasas 11824 (COL, F). Risaralda:
Municipio de Pereira, vicinity of La Laguna de Ottn, 4000-4300 m, 3.ii.1980 (fl),
Diaz, Valencia & Jaromillo 1738 (COL). Santandér: Paramo de Santurban, Laguna
de Cunta, 3880 m, 21.1.1927 (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
Map 10 20. H. laricifolium: typical form @, ‘laricoides’ O; 21. H. martense A.
49
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, 11.11.1944 (fl), Steyermark 55501 (F). Mérida: Paramo de Mucuchies,
x.1865 (fl), Moritz 1166 (BM, K, P, S); Sierra Nevada de Mérida, between La
Canada del Espejo and La Canada de Los Anteojos, Torre de Befio, 4000-4130 m,
22.1.1973 (fl), Cuatrecasas, Ruiz-Terdn & Lépez-Figueras 28570 (BM, US); Paramo
de Conejos, c. 8 km NE. of Mérida, 3800-4100 m, 27.xii.1967, Wessels Boer 2135
(NY, U, VEN). Tachira: Paramo de La Negra, 3000 m, 13.xi.1939 (fl), Burrus &
Miiller 1019a (VEN). Trujillo: El Paramito, Jag6 to La Morita, hacia Tufiame, over
3000 m, vili.1958 (fl), Aristeguiéta & 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.1969 (fl),
B. & C. Maguire 61696 (BM, NY). Bolivar: Cordillera Occidental, Gualicon Loma,
3240 m, 7.x.1943, Acosta Solis 6263 (F). Cafiar: Biblian to Canar, north of Biblian, c.
3400 m, 23.iv.1968 (fl), Harling, Storm & Strém 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, 3700 m, 3.vii.1939 (fl), Asplund 7516
(K, P, S). El Oro: NE. Zaruma, Tioloma, Hac. Ambocas, 3100 m, 30.viii.1947,
Espinosa 2034 (NY). Imbabura: Cotacache to Apuela, 21 km (Intec valley), 3300 m,
11.viii.1976 (fl), Dllgaard & Balslev 8687 (AAU, BM, MO, U). Loja: 10 km south of
Loja, 2800 m, 1-3.viii.1959 (fl), Harling 6239 (S). Napo: Cerro Antisana, Los
Corrales, near Papallacta, 3900 m, 21.vii.1960 (fl), Grubb et al. 218 (K, NY).
Pichincha: Mount Guamani, 3600-3800 m, 15.ix.1939 (fl), Asplund 8757 (K, P, S).
Tungurahua: between Ambato and Banos, 10.i.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. 1964 (fl), Hutchinson & Wright 4392 (F, K, MO, P, S). Cajamarca: 28 km N. of
Cajamarca towards Pedernal, 12 km W. to Hacienda Granja Porcon (SIPA), 3260
m, 6.vi.1963 (fl), D. & V. Ugent 5463 (GH, K, WIS). Huanuco: Mito, c. 2700 m,
8—22.ii.1922 (fl & fr), Macbride & Featherstone 1610 (F, S). La Libertad: Huama-
cucho, Pullac to Parcay, 69 km, above Rio Chuagén, 3920 m, 7.viii.1964 (fl),
Hutchinson, Wright & Straw 6188 (F, K, MO). Piura: Huancabamba, distr. Sondor,
2300 m, 21.vii.1975 (fl), Sagdastegui, Cabanillas & Dias 8214 (MO). San Martin:
Huallaga, valley of Rio Apisoncho 30 km above Jucusbamba, 3300 m, 5.viii.1965
(fl), Hamilton & Holligan 1090 (K, S).
H. laricifolium is closely related to H. thuyoides, and one collection from Cauca
(Fernandez Pérez 7159) is intermediate in some characters. Nearly always, however,
H. laricifolium can be distinguished by the narrow rounded leaves with impressed
midrib and straight margin.
H. laricifolium is very variable, but the variation appears to be continuous and not
amenable to classification. From northern Ecuador and adjacent Colombia, where
the leaves are relatively broad, there are clines north and south towards shorter,
narrower, more densely imbricate leaves and smaller flowers with narrower, more
acute sepals:
(i) North-eastward along the Cordillera Central to Venezuela; the extreme form
from Boyaca north-eastward looks very different, with young shoots densely clothed
with appressed leaves (‘H. laricoides’). Both the typical form and ‘H. laricoides’
occur in Boyaca and some parts of Cundinamarca, the latter favouring damper
habitats. In other parts of Cundinamarca (e.g. Paramo de Guasca) and in Caldas,
Risaralda, and Tolima, however, there is a continuous series of intermediate forms
that does not allow the recognition even of subspecies.
(ii) Southward through Ecuador to central Peru (Hudnuco, Ancash) the leaves
become gradually narrower and smaller, but not markedly imbricate, except for
some populations from northern Peru.
THE GENUS HYPERICUM L.
21. Hypericum martense N. Robson, sp. nov.
H. laricifolio Juss. affinis, sed habitu multo humiliori caespitosa, 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-canaliculate, cucullate, midrib not
impressed, margin narrowly hyaline, concolorous, both surfaces dull, glaucous,
subcoriaceous?; apex subacute to obtuse, base parallel, not clasping, pairs united to
form very narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense,
large, impressed beneath. Inflorescence 1-flowered, with pseudo-dichotomous bran-
ches from node below; pedicel 3—4 mm long, slender; upper leaves not transitional.
Flowers 8—10 mm in diam., obconic (?). Sepals 4—5 x 0-8—1-3 mm, narrowly oblong,
acute, incurved above; veins 5, unbranched, all prominent; glands mostly linear but
striiform to punctiform near hyaline margin. Petals 6-8 x 2 mmc. 1-5 X sepals,
oblong-obovate; apiculus acute; glands few, punctiform. Stamens c. 25, longest 4—5
mm long, c. 0-65 X petals. Ovary c. 1 x 0-5 mm, ellipsoid; styles 3, 2—2-5 mm long, c.
2-2-5 X ovary, outcurving to erect; stigmas broadly capitate. Capsule 4-5 x 2-2-5
mm, narrowly ellipsoid, obtuse, exceeding sepals. Seeds 1-2 mm long, ecarinate;
testa finely scalariform-reticulate.
In rocky paramo, muddy places in high-rainfall areas; 3000-3900 m.
Colombia (Magdalena); confined to the Sierra de Santa Marta. Map 10 (p. 49).
COLOMBIA. Magdalena: Sierra de Santa Marta, SE. slopes, Hoya del Rio
Donachui, Meollaca, c. 3400 m, 8.x.1959, Cuatrecasas & Romero Castaneda 24644
(US); above San Miguel, c. 3000 m, vii.1932, Seifriz 421 (US).
H. martense is a high-altitude derivative of the ‘laricoides’ form of H. 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
Pl. Hartwegiana: 127 (1843) [‘hartwegi’]; Triana & Planchon in Anals Sci. nat. (Bot.)
IV, 18: 295 (1862) pro parte, quoad typum; R. Keller in Engler & Prantl, Nat.
Pflanzenfam. 3 (6): 214 (1893), in Bull. Herb. Boissier II, 8: 177 (1908), in Engler
& Prantl, op. cit. 2nd ed. 21: 181 (1925) pro parte, excl. pl. Colomb. Type:
Ecuador, Loja, mountains near Loja, Chiquiribamba, vii.1841 (fl), Hartweg 721
(K!, holotype; BM!, K!, P!, W!, isotypes).
Shrub 0-3-0-9 m tall, erect, with branches strict to spreading, lateral, short (flower-
ing) and long (extension) or rarely pseudo-dichotomous. Stems dark red-brown,
4-lined and ancipitous when young, eventually terete, without epidermal emerg-
ences, cortex exfoliating in strips; internodes 5-7 mm long. Leaves sessile, erect to
ascending, closely imbricate, tetrastichous, persistent; lamina 5-7 x 2-3 mm,
elliptic to oblong or obovate, plane, apically subconcave, not carinate but with
midrib + incrassate distally and subimpressed proximally beneath, margin narrowly
hyaline, not glaucous, subcoriaceous; apex apiculate-obtuse to rounded, slightly
cucullate, base cuneate to parallel, scarcely clasping, pairs united to form very
narrow interfoliar ridge; basal vein 1, apparently unbranched; laminar glands quite
51
32
O HEE
O Hh =
NORMAN K. B. ROBSON
dense, not impressed or prominent, visible on both sides. /nflorescence 1-flowered,
terminal and on short lateral branches; pedicel 1-2 mm long; upper leaves foliose.
Flowers 15—20 mm in diam., stellate. Sepals 4-6 x 1-8-2-3 mm, ovate to oblong-
elliptic, acute to subacute; veins c. 5, unbranched (?), midrib distally incrassate;
glands linear. Petals bright (?) yellow, 8-11 x 4-5 mm, 2-2-5 x sepals, obovate;
apiculus acute; glands striiform and punctiform. Stamens 45-50, longest 6-7 mm
long, c. 0-5 X petals. Ovary 2-2-5 x c.1-4 mm, ovoid-ellipsoid; styles 3, 5-6 mm
long, c. 2-5 X ovary, Outcurving; stigmas small to subcapitate. Capsule and seeds not
seen.
Habitat and phenology not recorded.
Ecuador (Loja). Known from only one collection. Map 9 (p. 46).
ECUADOR. Loja: see type.
H. hartwegii appears to be a very local relict species, being mostly closely related to
H. lycopodioides (from east-central Colombia), which has spreading, incurved-
conduplicate leaves with margin sometimes undulate, oblique-obovate petals, and
more numerous stamens. See also 23 H. maguirei.
23. Hypericum maguirei N. Robson, sp. nov.
H. hartwegii Bentham affinis, sed foliis minoribus, dense imbricatis vaide cucullatis
costa media subtus impressa, floribus minoribus, differt. Type: Ecuador, Azuay,
5km past San Miguel on road to Cuenca, 3350m, 15.x.1969 (fl), B. & C. Maguire
61707 (BM!, holotype; NY!, isotype).
Shrub 0-3-1 m tall, erect or spreading; branches strict or ascending, lateral, short
(flowering) and long (extension), narrowly spiciform to pyramidal, sometimes
secund. Stems dark red-brown, 4—6 lined when young, eventually terete, without
epidermal emergences, cortex exfoliating in strips or irregular flakes; internodes 1-3
mm, long. Leaves sessile, suberect to erect, + densely imbricate, tetrastichous,
persistent; lamina 2—4 x 0-4—-1-2 mm, + narrowly oblong to narrowly elliptic or
linear, incurved, midrib impressed beneath, pale lucent green with margin narrowly
hyaline and plane, not glaucous, coriaceous; apex obtuse to rounded, markedly
cucullate, base narrowly cuneate to parallel-sided, not or scarcely clasping, pairs
united to form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands
dense or in one row beneath, not impressed or prominent, sparser above. /nflores-
cence 1-flowered, terminal and on short lateral branches; pedicel 1-3 mm long;
upper leaves foliose. Flowers (S—)8—15 mm in diam., stellate (to obconic ?). Sepals
2-5 xX 0-7-7-1 mm, acute, incurved-cucullate, margin hyaline; veins 3-5, unbran-
ched, midrib prominent beneath; glands linear, distally punctiform. Petals golden to
orange-yellow, 3-8 xX 1-5-4 mm, 1-5-2 xX 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 Ofia y Rancho Ovejero, 2704—2800 m, 1-2.viii.1959 (fl), Barclay &
Juajibioy 8452 (NY). El Oro: between Curtincapa and Guagra Uma, 13 km NE. of
Curtincapa, 2850 m, 16.viii.1943 (fl), Steyermark 53912 (F). Loja: about Ofia, 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 Santandér, 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
ao
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 %; c—g x 2). (b) Cuatrecasas & Rodriguez 27789; others
Steyermark 57378.
54
NORMAN K. B. ROBSON
not deflexed, persistent until cortex is shed; lamina 10-18 x 4—8 mm, elliptic to
oblanceolate-spathulate, plane, slightly cucullate, carinate, concolorous or paler
beneath, not glaucous, chartaceous; apex acute to obtuse-apiculate, base angustate,
sheathing, pairs united to form very narrow interfoliar ridge; basal veins 5—7, parallel
or almost so, branching distally, with tertiary reticulation sometimes visible beneath;
laminar glands dense above, sparse or absent beneath, slightly prominent. /nfloresc-
ence 1-flowered, sometimes with several short flowering branches crowded together,
usually with pseudo-dichotomous branches from Ist or 2nd node below; pedicel up to
5 mm long; upper leaves transitional. Flowers c. 35 mm in diam., stellate. Sepals 9-11
x 3-5-5 mm, broadly oblong to obovate-spathulate, acute to obtuse; veins 11-19,
dichotomising distally, with midrib scarcely prominent beneath; glands punctiform
in upper third. Petals rich yellow, 16-22 x 12-14 mm, c. 2 X sepals, obovate-
oblanceolate; apiculus acute; glands punctiform distally. Stamens c. 80, longest 9-11
mm long, c. 0-5 x petals. Ovary 4-5 x 2-3 mm, broadly ellipsoid to subglobose;
styles 3, S—9 mm long, 1-5-2 X ovary, erect, free, outcurved below apex; stigmas
slightly enlarged to broadly capitate. Capsule c. 7 X 5mm, broadly ellipsoid, shorter
than sepals. Seeds not seen.
On rocky slopes of limestone outcrops; 3000—3800 m.
Eastern Colombia (Boyaca, Santandér, Norte de Santandér) and adjacent Vene-
zuela (Tachira). Map 11.
COLOMBIA. Boyaca: Paramo de Chita, cabeceras de Rio Casanare, 3080 m,
19. vi. 1972 (fl), Cleef 4743 (BM, U). Norte de Santandér (see also type): Bucaraman-
ga, Cumbre of Paramo Mortifo, Cucuta road, 3300 m, v.1948 (fl), Sandeman 6077
THE GENUS HYPERICUM L.
(K). Santandér: Paramo de Santurban, c. 3000 m, 27.viii.1948 (fl & fr), Barkley &
Araque 188155 (F, COL).
VENEZUELA. Tachira: Paramo de Tama, near Colombo-Venezuelan bound-
ary, 3045-3475 m, 15.vii.1944 (fl). Steyermark 57378 (F, NY); NNE. slopes of Pata
de Judio, 2500-3100 m, 19.x.1978 (fi & 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 witha
sheathing narrowed base.
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, Santandér, 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 bot. Club 56: 106. (1929) pro parte, quoad
Killip & Smith 15616 pro parte.
Shrub 0-15-1-5 m tall, erect, with branches strict, mostly pseudo-dichotomous,
sometimes lateral. Stems yellow-brown, 4-lined when young, eventually terete,
without epidermal emergences, cortex flaking irregularly; internodes 2-3 mm long.
Leaves sessile, outcurved-ascending to imbricate-erect, tetrastichous, deciduous
above base before fading; lamina 7-10 x 3-6 mm, oblanceolate to obovate,
incurved, not cucullate or carinate, concolorous, not glaucous, subcoriaceous to
coriaceous; apex acute or apiculate to obtuse, base angustate, sheathing, pairs united
(at least when young) to form narrow interfoliar ridge; basal veins 5—7, flabellate,
branching distally (sometimes obscurely), reticulation obscure beneath; laminar
glands dense, not or slightly prominent, visible beneath; marginal glands usually
secreting viscous clear or white resin. Inflorescence 1-flowered usually with pseudo-
dichotomous branches from 1st to 3rd node below and often with 1 or 2 lateral
branches immediately below these, forming flower cluster when young; pedicel 3—6
mm long; upper leaves transitional. Flowers 20-40 mm in diam., stellate. Sepals
6-10 x 3-5 mm, elliptic to oblanceolate, acute to apiculate or obtuse; veins 9-15,
sometimes dichotomising distally, with midrib scarcely prominent beneath; glands
punctiform in upper third to two-thirds. Petals rich yellow, (always?) tinged red
beneath, 12—20 x 10-11 mm, c. 2 X sepals, oblanceolate to obovate; apiculus obtuse
to rounded; glands distally punctiform. Stamens 80-100, longest 6-10 mm long, c.
0:5 x petals. Ovary 3-4 x 2-3 mm, broadly ellipsoid to ovoid; styles 3, 4-6 mm long,
1-25-1-5 x ovary, free, suberect to spreading; stigmas broadly capitate. Capsule 5—6
x 5-6 mm, subglobose, shorter than sepals. Seeds not seen.
In open paramo on fine talus or grassy slopes; 3200-3900 m.
Colombia (Norte de Santandér, Santandér). Apparently confined to the Paramo del
Almorzadero. Map 9 (p. 46).
COLOMBIA. Norte de Santandér: Valley of rio Chitaga, 20 km SE. of Chitaga,
3475 m, 24.ix.1944 (fl & fr), St. John 207784 (COL, NY). Santandér: Péramo del
Almorzadero, 3600-3800 m, 28.xi.1941 (fl), Cuatrecasas 13495 (COL, GH, NY, P);
Péramo del Almorzadero, between Chitag4 and Cerrito, 3900 m, 31.xii.1959-
1.i1.1960 (fl), Barclay & Juajibioy 10389 (NY); Peralonso, Platera, 3400 m, 20.ix.1969
(fl), Cuatrecasas & Rodriguez 27828 (BM, COL, US); Peralonso, 3200 m,
19.vii.1940 (fr), Cuatrecasas & Garcia Barriga 9887 (COL).
H. gleasonii is a relict species intermediate between H. magniflorum and H.
mexicanum. It has larger flowers than H. mexicanum, and these are solitary, but it
shares with that species the deciduous leaves with viscous secretions. In H. gleasonii
the leaves are smaller than those of H. 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 (1776), Amoen. Acad. 8: 322, t.8, f.2 (1785); L. fil., Suppl. PI.:
345 (1781); Lam., Encyl. méth. (Bot.) 4: 169 (1797); Gleason in Bull. Torrey bot.
Club 56: 103 (1929). Type: Colombia [Cundinamarca], Mutis in Herb. Linn. 943:
31 (LINN!, holotype; BM!, isotype).
Hypericum ? mexicanum sensu Choisy, Prodr. monogr. fam. Hypéric.: 60 (1821).
Hypericum mutisianum Kunth in Humboldt, Bonpland & Kunth, Nov. Gen. et Sp.
PI. 5: 185 (1822), nom. illegit.; R. Kellerin Bull. Herb. Boissier I, 8: 178 (1908), in
Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925); Knuth in Reprium Sp.
nov. Regni veg., Beih. 43: 483 (1927). Type: Colombia, Cundinamarca, near Santa
Fé de Bogota, 2457 m, 1805 (fl), Humboldt & Bonpland s.n. (P-HUM!, holotype;
B-WILLD, isotype);
Brathys mutisiana (Kunth) Spach, Hist. nat. vég. Phan. 5: 447 (1836); in Annls Sci.
nat. (Bot.) II, 5: 366 (1836).
Hypericum resinosum Bentham, Pl. Hartweg.: 165 (1845); R. Keller in Bull. Herb.
Boissier I1, 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. pl. 1: 59 (1891). Type: Vene-
zuela, Mérida, Silla de Caracas, Kuntze s.n. (NY, holotype).
Shrub or shrublet 0-15—1-5(—2) m tall, erect or decumbent and sometimes rooting at
the base, with branches strict, pseudo-dichotomous and sometimes lateral especially
from short upper internodes. Stems yellow- to orange-brown, 4-lined when young,
eventually terete, cortex flaking irregularly; internodes 2—8 mm long. Leaves sessile,
outcurving-ascending to imbricate-erect, tetrastichous, deciduous above base usu-
ally without withering; lamina 7-17 x 3-10 mm, elliptic or oblanceolate to obovate,
plane to subconduplicate, not or scarcely cucullate, not carinate, concolorous, not
glaucous, coriaceous; apex obtuse or rounded, base cuneate to angustate, +
sheathing, pairs united to form narrow interfoliar ridge; basal veins 5—7, flabellate,
branching and anastomosing distally, reticulation clearly to obscurely visible be-
neath; laminar glands dense to sparse, usually visible and sometimes slightly
prominent beneath; marginal glands secreting viscous clear or white resin. Inflores-
cence 1—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 (Mérida, Tachira) and north-eastern to central Colombia (Norte
de Santandér to Valle de Cauca). Map 12.
COLOMBIA. Antioquia: entre Rio Negro y Sta Helena, 20650 m, vii.1944 (fi),
Garcia Barriga. 11094 (COL). Boyaca: Sierra Nevada del Cocuy, Quebrada de San
THE GENUS HYPERICUM L.
Map 12 26. H. mexicanum ®; 28. H. prietoi O; 29. H. cassiopiforme @.
Paulino proximo Alto Ritacuva, c. 3650, 3.v.1959 (fl), Barclay & Juajibioy 7525
(COL, MO, NY); Paramo de Gina, 3200 m, 17.vii.1940 (fl), Cuatrecasas & Garcia
Barriga. 9783 (COL). Cundinamarca: Paramo de Guasca, 2840 m, 15.xii.1938 (fl),
Balls 5700 (BM, COL, K); Sabana de Bogota, Municipio de Tenjo, Vereda de la
Punta, 2800-2670 m, 29.i.1966 (fl & fr), Cuatrecasas & Jaramillo 26755 (BM, COL,
US); Cerro de Monserrate, 3000 m, 10.iii.1966 (fl), Duque-Jaramillo 2877 (COL).
Meta: Macizo de Sumapaz, alredores de la Laguna La Guitarra, 3370-3400 m,
5.vii.1981, Diaz P. 2485 (COL). Norte de Santandér: Paramo de Fontib6n, 2700 m,
27.vii.1940 (fl), Cuatrecasas & Garcia Barriga. 10088 (NY, US). Santandér: 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. Mérida: 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 of H. mexicanum with H. platyphyllum,
which has penninerved (not flabellate-nerved) leaves and smaller stigmas and is
treated here as a subspecies of H. phellos.
ay
NORMAN K. B. ROBSON
27. Hypericum stuebelii Hieron.
in Bot. Jahrb. 21: 321 (1895); R. Keller in Bull. Herb. Boissier I1, 8: 177 (1908), in
Engler & Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925) Type: Peru, Amazonas,
Chachapoyas, between Pacasmayo and Moyobamba, near Mojon Cruz, 3300 m,
iv.—vi.1875? (fl), Sttibel Peru 27 (B+; 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. Jn-
florescence 1-flowered, usually with pseudo-dichotomous branches from node be-
low; pedicel 4—6 mm long; upper leaves not transitional. Flowers 25—40 mm in diam.,
stellate. Sepals 10-12 x (2—)3—6 mm, elliptic-ovate to lanceolate, obtuse to acute,
cucullate; veins 7-11, sometimes dichotomising distally, prominent beneath; glands
linear, punctiform in distal third. Petals bright? yellow, 15S—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.1962 (fl), Wurdack 1353 (F, GH, K, NY, P, S, US); La Jalca
near Chachapoyas, 2700-3300 m, i.1930 (fl), Williams 7585 (F, NY); Chachapoyas,
Pass of Piscohuafuna, 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 proximali subtus
costa media impressa, floribus minoribus, differt. Type: Ecuador, Azuay, Eastern
Cordillera, between Ona and the rio Yacuambi, 3000-3360 m, 10-19.ix.1945 (fr),
Prieto P-302 (NY!, holotype).
Shrub c. 0-3 m tall, erect; branches very strict, pseudo-dichotomous and lateral.
Stems reddish-brown, 4—6-lined when young, eventually terete, cortex exfoliating in
irregular flakes; internodes c. 2 mm long. Leaves sessile, very closely imbricate,
markedly tetrastichous, laterally compressed distally, not becoming outcurved,
deciduous above the base before fading; lamina 7-8 x 1-5-2 mm, oblanceolate,
distally obtusely carinate and cucullate, midrib impressed proximally beneath,
margin very narrowly hyaline, concolorous, lustrous yellow-green, not glaucous,
coriaceous; apex acute, base parallel-sided, scarcely sheathing, pairs united to form
shallow to rather deep (to 1-5 mm) interfoliar ridge; basal vein 1, unbranched;
THE GENUS HYPERICUM L.
laminar glands dense, not visible beneath. Inflorescence 1-flowered, with pseudo-
dichotomous branches from node below; pedicel 2—2-5 mm long; upper leaves not
transitional. Flowers stellate? Sepals 7-8 x 2 mm, lanceolate, acuminate, compress-
ed-cucullate; veins 3—5, unbranched, prominent beneath; glands linear. Petals and
stamens not seen. Ovary with styles 3, c. 4 mm long, suberect; stigma narrowly
capitate. Capsule 5 X 2-5 mm, broadly ellipsoid, shorter than sepals. Seeds 1-2 mm
long, ecarinate; testa finely scalariform.
On the crest of the Cordillera; 3000-3360 m.
Ecuador (Azuay). Map 12 (p. 57).
ECUADOR. Azuay: Eastern Cordillera between Ona and the rio Yacuambi,
3000-3360 m, 10—19.ix.1945 (fr), Prieto P-302 (NY).
H. prietoi, which has been collected once only, resembles H. stuebelii but has
smaller, laterally compressed, and markedly tetrastichous leaves and stricter
branches, some of which are lateral.
29. Hypericum cassiopiforme N. Robson, sp. nov.
H. stuebelii Hieron. affinis, sed ramis basalibus lateralibusque strictioribus, foliis
minoribus, oblanceolatis vel obovato-oblanceolatis, floribus minoribus, staminis
paucioribus, stylis brevioribus, differt. Type: Peru, Amazonas, Chachapoyas, 1-5
km W. of Molinopampa, 2400-2450 m, 18.vii.1962 (fl), Wurdack 1355 (US!,
holotype).
Shrub c. 0-3 m tall, erect, with branches strict, basal and lateral. Stems reddish-
brown, 4-lined when young, eventually terete, cortex exfoliating in irregular flakes;
internodes 2—4 mm long. Leaves sessile, imbricate-tetrastichous but not usually
laterally compressed distally, becoming outcurved, deciduous above the base before
fading; lamina 7-8 x 2-5-3 mm, oblanceolate, incurved-cucullate but scarcely
carinate, midrib slightly prominent to scarcely impressed beneath, margin narrowly,
hyaline otherwise concolorous, lustrous yellowish-green, not glaucous, sub-
coriaceous; apex subacute, base cuneate, broadening below, scarcely sheathing,
pairs united to form narrow interfoliar ridge; basal vein 1 with 2 obscure near-basal
branches; laminar glands dense, more visible above than beneath. Inflorescence
1-flowered; pedicel c. 3-5 mm long; upper leaves not transitional. Flowers c. 25 mm in
diam., stellate. Sepals 8 x 1-8—2-3 mm, elliptic to oblong, acute, incurved-cucullate,
reddish; veins 5, unbranched, prominent beneath; glands linear, distally punctiform.
Petals bright yellow tinged red, c. 14 x 9mm, c. 1-7 X sepals, obovate-oblanceolate;
apiculus acute; glands linear. Stamens c. 50, longest c. 10-11 mm long, c. 0-5 x
petals. Ovary c. 4 X 1:7 mm, ellipsoid; styles 3, c. 5 mm long, c. 1-2 X ovary, sub-
erect ?; stigmas narrow. Capsule and seeds not seen.
In the Jalca zone; 2400-2450 m.
Peru (Amazonas). Map 12 (p. 57).
PERU. Amazonas: 1—5 km W. of Molinopampa, 2400-2450 m, 18.vii.1962 (fl),
Wurdack 1355 (US).
H. cassiopiforme, like H. prietoi, is based on a single collection and appears to be
related directly to H. stuebelii. It is smaller, with smaller leaves and flowers and more
slender stems, and is intermediate in form between H. stuebelii and H. decandrum. It
differs from the latter by the broader leaves, the larger flowers with broader sepals
and the longer styles with narrow stigmas.
30. Hypericum decandrum Turcz.
in Bull. Soc. Nat. Moscou 31 (1): 389 (1858). Type: Ecuador [Pichincha (?)], Quito,
[18657] (fl), Jameson 62 (LE, holotype; BM!, K!, W!, isotypes).
Hypericum gnidioides var. polytrichoides R. Keller in Bull. Herb. Boissier 11, 8: 183
(1908). Type: Ecuador, Chimborazo, in pascuis montis Titaicin, 3900 m, xi.1859
(fr), Spruce 5599 (G!, holotype, F!, photograph; BM!, C!, GH!, K!, NY!, P!, W!,
isotypes).
\
Y
ag
NORMAN K. B. ROBSON
Icon: Fig. 10.
Shrub or shrublet (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 xX 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). Caniar: Rivera NE. of
Pindilig, 2900 m, 13.xii.1980 (fi & fr), Holm-Nielsen, Jaramillo & Coello 29222
(AAU). Carchi: Nudo de Boliche, Voladero, 3800 m, 12.vi.1939 (fl & fr), Penland
900 (F, GH, NY). Chimborazo: Mt. Titaictin, 3900 m, xi.1858 (fl), Spruce 5599 (BM,
C, GH, K, NY, P, W). Loja: N. of Saraguro, 3045 m, 4.viii.1943 (fr), Steyermark
53743 (F, NY). Morona-Santiago: Paramo de Hatanga, Km 36 on road Sigsig-
Gualaquiza, E. of pass, 3300 m, 14.xii.1980 (fl), Holm-Nielsen, Jaramillo & Coello
29386 (AAU). Napo: Valle de Chalupas near junction of Rio Chalupas and Rio
Aguas Buenas, 3200 m, 20.i.1980 (fl), Holm-Nielsen 21014 (AAU). Pichincha?:
Quito, 1865? (fl), Jameson 62 (BM, K, W). Tungurahua: Cordillera de Llanganates,
near junction of rio Golpe and rio Sangarinas (Desaguadero), 26.xi.1939 (fl & fr),
Asplund 9989 (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 Santandér, Prov. de Ocafia, 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 I, 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 4%; 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 1-5(—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
NORMAN K. B. ROBSON
obscure beneath. /nflorescence 1-flowered, with pseudo-dichotomous branches from
one or two nodes below; pedicel 4—12 mm long, incrassate upwards; upper leaves not
transitional. Flowers 18-35 mm in diam., stellate to subcyathiform. Sepals (7)10—14
x (2)3-8 mm, elliptic-subcircular to elliptic-oblong or rarely oblong, acute to
subacute; veins (5)7—11, dichotomising distally, with midrib not or slightly promin-
ent; glands linear, distally punctiform. Petals bright yellow, 10-22 x 4-10 mm, c.
1-3-1-5 X sepals, obovate; apiculus acute; glands linear, distally punctiform. Sta-
mens c. 40-90, longest 5—10 mm long, c. 0-5 x petals. Ovary (2-5)3—4 x 2-2-5 mm,
ovoid-ellipsoid to subglobose; styles 3—4, (2)3—7 mm long, 1—1-75 x ovary, outcurv-
ing to diverging, stout; stigmas scarcely to broadly capitate. Capsule 5—8 x 4-6 mm,
cylindric-ellipsoid, equalling sepals. Seeds c. 1-2 mm long, ecarinate; testa finely
scalariform.
In open paramo; 3000-4300 m.
Colombia (César/Norte de Santandér 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 Playén, Agua
Tendida, 3100 m, 10.vi.1973 (fl), Cleef 10174 (U). Boyaca: Sierra Nevada del Cocuy,
near El Play6én, c. 3800 m, 10.ix.1957 (fl), Grubb, Curry & Fernandez-Perez 777
(COL, K, US); Paramo de Chita, cabeceras de Rio Casanare, Km 93 de la carretera a
La Punta, 3080 m, 19.vii.1972 (fl), Cleef 4744 (BM, U); Pea de Arnical, N. de Vado
Hondo, 3600 m, 6.iv.1973 (fl), Cleef 9418 (U). César/Norte de Santandér: 20 kmalS.
Map 13 31. H. pimeleoides @; 32. H. magdalenicum A; 33. H. valleanum A; 34. H. sprucei @; 37. H.
wurdackii O.
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 Santandér: Paramo de Las
Jurisdicciones, SW. of Ocania, 3000-3500 m, 8.vii.1984 (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.
32. Hypericum magdalenicum N. Robson, sp. nov.
H. pimeleoidei Planchon & Linden ex Triana & Planchon affinis, 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—-)0-7-1-5 m tall, erect, with branches strict, pseudo-dichotomous and
sometimes with 2—3 laterals closely beneath flower. Stems orange-brown, 4-lined and
compressed when young, the subfoliar ridges broad, soon terete, cortex exfoliating
irregularly; internodes 2—4 mm long. Leaves sessile, densely imbricate, erect or
outcurving, markedly tetrastichous, deciduous above base without fading; lamina
8-18 x 2-6-2 mm, broadly to rather narrowly elliptic or oblanceolate, plane to
incurved-conduplicate, not or slightly cucullate, midrib plane or slightly impressed
beneath, margin not distinct, epidermis sometimes undulate, concolorous, some-
times glaucous, + thinly to thickly coriaceous; apex acute, base narrowly cuneate,
scarcely sheathing, pairs united to form narrow interfoliar ridge; basal vein 1, with c.
3—4 pairs of ascending lateral branches, sometimes branching distally, not prominent
or impressed beneath, tertiary reticulation not visible; laminar glands + dense,
sometimes visible beneath. Inflorescence 1-flowered, with pseudo-dichotomous
branches from node below; pedicel 4-5 mm long, not incrassate upwards; upper
leaves not transitional. Flowers 20-30 mm in diam., stellate. Sepals (7—)9-11 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-)9-11 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 Mérida). In the Sierra Nevada de
Santa Marta and the Sierra de Perija, with one probably conspecific record from the
Paramo del Molino. Map 13.
COLOMBIA. Magdalena: Sierra Nevada de Santa Marta, alredores de cabeceras
64
NORMAN K. B. ROBSON
de Rio Sevilla, 3320-3410 m, 22.1.1959 (fl & fr), Barclay & Juajibioy 6620 (COL,
MO, NY); Sierra Nevada de Santa Marta, valley of Rio Yebosimeina, 3500—4000 m,
22.v.1977 (fl & fr), Starker White & Alverson 547 (COL, MO, NY); Sierra de Perija,
E. of Manaure, quebrada de Floridablanca, 2700-2800 m, 9—12.xi.1959, Cuatre-
casas & Romero Castaneda 25171 (COL).
VENEZUELA. Mérida: Péramo del Molino, 2600 m, 19.i.1922 (fl), Jahn 940
(VEN). Zulia: Perija, Cerro Fetari, iv.1952 (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 affinis, sed foliis latioribus, nec profunde incurvatis nec
cucullatis, manifeste glanduloso-punctatis, pedicello 3—4 mm longo, stylis crassius-
culis, differt. Type: Colombia, Valle de Cauca, Los Farallones, filo de la cordillera,
matorrales de paramo en el cerro La Torre, c. 3750 m, 10.x.1944 (fl), Cuatrecasas
177861 (F!, holotype & isotype).
Shrub c. 1 m tall, erect, with branches strict, pseudo-dichotomous and lateral. Stems
orange-brown, compressed? when young, soon subterete to terete, cortex exfoliat-
ing irregularly; internodes 1-2 mm long. Leaves sessile, subimbricate, narrowly
spreading and twisting, subtetrastichous, deciduous above base without fading;
lamina 11-17 xX 2-3 mm, narrowly elliptic, shallowly incurved, not cucullate, midrib
impressed, margin very narrowly hyaline, concolorous, not glaucous, coriaceous;
apex sharply acute, base angustate, not sheathing, pairs united to form very narrow
interfoliar ridge; basal vein 1, unbranched; laminar glands dense, large, visible
beneath. Inflorescence 1-flowered, with pseudo-dichotomous branches from node
below; pedicel stout, 3—4 mm long; upper leaves not transitional. Flowers c. 20 mm in
diam.?, stellate. Sepals 7-10 x 2-2-8 mm, narrowly elliptic to lanceolate, acute to
acuminate, incurved above, margin narrowly hyaline, veins 5, the outer branched,
midrib not or scarcely prominent; glands linear, distally punctiform. Petals bright (?)
yellow, 10-12 X 5-6 mm, c. 1:2-1-4 X sepals, obovate; apiculus sharply acute;
glands striiform to punctiform. Stamens c. 60 ?, longest 5-6 mm long, c. 0-5 x petals.
Ovary c.2 X 1:5mm, ellipsoid; styles 3, c. 3mm long, c. 1-5 x ovary, stout, suberect;
stigmas narrow. Capsule and seeds not seen.
In paramo; 3750 m.
Colombia (Valle de Cauca). Known only from the type collection. Map 13 (p. 62).
COLOMBIA. Valle de Cauca: Cordillera Occidental, Los Farallones, cerro La
Torre, c. 3750 m, 10.x.1944 (fl), Cuatrecasas 17861 (F).
H. valleanum is intermediate in form and distribution between the more primitive
form of H. magdalenicum (with broad punctate leaves) and H. sprucei, differing
from the former by its shorter styles and narrower leaves, and from the latter by its
broader, thicker, and densely punctate not cucullate leaves and its stout styles and
generally stronger habit. It appears to be a relict population.
THE GENUS HYPERICUM L.
34. Hypericum sprucei N. Robson, sp. nov.
H. aciculari Kunth 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.1857
(fl), Spruce 5110 (BM!, holotype; c!, GH!, K!, P!, S!, isotypes).
Hypericum struthiolifolium sensu R. Keller in Bull. Herb. Boissier II, 8: 180 (1908)
[‘struthiolaefolium’|; 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
linear, distally punctiform. Petals bright to deep yellow, 10-15 x 5-7 mm, c. 1-4 x
sepals, oblong-oblanceolate; apiculus sharply acute to apiculate; glands striiform to
punctiform, sparse. Stamens 60—75, longest 5—9 mm long, 0-5-0-6 x petals. Ovary
2-3-5 x 1-5-2 mm, ellipsoid-subglobose; styles 3(4), (2:5)3—4-5 mm long, (1-3)1-5
—2 X ovary, outcurving, slender; stigmas narrowly or not capitate. Capsule c. 5 x 3
mm, broadly ellipsoid to subglobose, shorter than sepals. Seeds c. 0-8 mm long
(immature?), ecarinate; testa finely scalariform.
Open, dry or damp paramo; 2725-4000 m.
Ecuador (Carchi to Loja, apparently absent from Cotopaxi and Bolivar), Peru
(Piura). Map 13 (p. 62).
ECUADOR. Azuay: c. 10 km NE. of Sevilla de Oro, 3000—3200 m, 13.ix.1976
(fl), Dllgaard & Balslev 9449 (AAU, BM, MO); ‘Oriente’ border, Eastern Cordil-
lera between Ona and rio Yacuambi, 3000-3360 m, 10—19.ix.1945 (fl & fr), Prieto
P-297 (BM, NY). Canar: between Biblian and Camiar, c. 3350 m, 21.ix.1955 (fl),
Asplund 17665 (S). Carchi: Base of Volcan Chiles, km 34—36 on Tulcan — Maldonado
road, 3900-4050 m, 19.v.1973 (fl), Holm-Nielsen et al. 5888 (AAU, F, GB, MO, S,
U). Chimborazo: Atilio, c. 3700 m, 29.i.1968 (fl), Harling, Storm & Strém 6705
(GB). Imbabura: Ibarra to Mariano Acosta, at pass (Hacienda Yura Cruz), 3600
—3800 m, 10.viii.1976 (fl & fr), Dllgaard & 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
(Lim6n) road, 3000-3100 m, 20.ic.1967 (fl), Sparre 18778 (S). Napo: Cordillera de
Los Llanganati, Chihuila Sacho o Ainchilibi, c. 3750 m, 25—29.viii.1959, Barclay &
Juajibioy 8996 (MO, NY); Sucumbios ‘Playon de San Francisco’, Camino al Cerro el
Mirador, 3600-3800 m, 29.xii.1980 (fl), Jaramillo & Coelle 3903 (AAU, QCA).
Pichincha: 15-4 km west of Papallacta, 3980 m, 26.iii.1972, MacBryde & Dwyer 1185
(MO). Tungurahua: Llanganati Mts, Lake Aucacocha, 3700 m, vii.1969, Edwards
58(K); Paramo of Minza Chica, 3800 m, 4.iv.1939 (fl), Penland 307 (F, GH). Zamora
—Chinchipe: Loja to Zamora, 2800 m, 18.ix.1961, Dodson & Thien 685 (BM).
PERU. Piura: above Huancabamba, 3000 m, iv.1912 (fl), Weberbauer 6133 (F,
GH, S).
H. sprucei provides a morphological and geographical link between H. valleanum
(Colombia) and three species in southern Ecuador and Peru (Spp. 35-37). From H.
65
:
66 NORMAN K. B. ROBSON
Cc
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 2; b x 2;
c-g X 3). A. Ollgaard & 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. Pl. 5: 190 (1822); Choisy in
DC., Prodr. syst. nat. regni veg. 1: 553 (1824); Jameson, Pl. Aequator. 1: 108
(1865); R. Keller in Bull. Herb. Boissier I1, 8: 176 (1908) ? (see below), in Engler
THE GENUS HYPERICUM L.
& Prantl, Nat. Pflanzenfam. 2nd ed. 21: 181 (1925). Type: Ecuador, Loja, locis
temporis prope Loxam (Regni Quitensi), 1908 m, viii.1805, Humboldt & Bon-
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. 16f. 2b).
H. struthiolifolium [var.] B sensu Choisy, Prodr. monogr. fam. Hypéric.: 59 (1821)
[‘struthiolaefolium’| pro parte, quoad spec. Peruv.
H. struthiolifolium [var.] B 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. vég. Phan. 5: 366 (1836), in Annis Sci.
nat. II, 5: 366 (1836).
H. laricifolium sensu Bentham, Pl. Hartweg.: 126 (1843-1844).
Shrub 0-3-2 m tall, erect and bushy to decumbent and slender, with branches strict to
ascending, lateral and frequently pseudo-dichotomous. Stems orange-brown, 4-lined
and + ancipitous when young, eventually terete, cortex exfoliating irregularly;
internodes 1-7 mm long. Leaves sessile, subimbricate or not, suberect, outcurving
and usually twisting, not or scarcely tetrastichous, deciduous above base without
fading; lamina 3-5-7 x 0-4—0-6 mm, very narrowly elliptic to linear-acicular,
incurved, slightly cucullate, midrib impressed beneath, margin very narrowly
hyaline, concolorous, not glaucous, subcoriaceous to chartaceous; apex acute, base
parallel-sided, not sheathing, scarcely broadening but pairs forming narrow inter-
foliar ridge; basal vein 1, unbranched; laminar glands dense to very sparse, visible
beneath. Inflorescence 1-flowered, terminal, with pseudo-dichotomous branches
from node below and terminating short unbranched lateral shoots; pedicel 1-5—2-5
mm long, not incrassate upwards; upper leaves not transitional. Flowers 10-17 mm
in diam., stellate. Sepals 4-7-5 x 0-7—1-5 mm, narrowly lanceolate, acute, incurved
above, margin hyaline, veins (3)5, midrib not prominent but whole sepal often
becoming ribbed; glands linear, distally punctiform. Petals yellow or apricot to
orange, sometimes tinged red, 6-10 x 3-7 mm, c. 1-5 X sepals, oblong-obovate,
apiculus sharply acute; glands striiform to punctiform. Stamens 30-55, longest 3-5
mm long, c. 0-5 X petals. Ovary 1-5—2-5 x 1-1-5 mm, ellipsoid-subglobose; styles 3,
2-5-3(4) mm long, 1-2-2 X ovary, outcurving-ascending; stigmas narrow. Capsule
3-5-4 x 2-5-3-5 mm, broadly ellipsoid, shorter than sepals. Seeds 1-2 mm long,
ecarinate; testa finely scalariform.
Dry, open scrub on slopes of paramo, dense moist forested slopes, and wet
sphagnum; 1800-3700 m.
Ecuador (Azuay, Loja), Peru (Piura, Amazonas). Map 14 (p. 68).
ECUADOR. Azuay: Km 85 on Pan-American Highway north of Loja, 2850—2950
m, 3.v.1973 (fl), Holm-Nielsen et al. 4808 (AAU, F, GB, S, U); Rio Collay south of
El Pan, Huagrarancha slopes, 2650-3290 m, 6.vii.1943 (fl), Steyermark 53372 (F);
*30-50 km S. of Cuenca, Paéramo de Tinajillas; 3300-3450, 17.ii1.1945, Camp
E-2229 (BM, NY);* Rancho Ovejero, c. 3000 m, 1-2.viii.1969 (fl). Barclay &
Juajibioy 8484 (NY). Loja: between La Toma and Loja, 1800-2600 m, 4.ix.1923 (fl),
Hitchcock 21431 (GH, NY, US);* 7 km from Loja on road to Zamora, 2350 m,
8.ix.1959 (fl), B. & C. Maguire 61667 (BM, NY);* W. slopes of Cerro Villonaco,
2200-2250 m, 10.iv.1974 (fl & fr), Harling & Anderson 13351 (GB, NY).
PERU. Piura: Huancabamba to Cuello del Indio, Mitopampa, 2650 m, 22.vii.1975
(fi), Sagdstegui, Cabanillas & Dios 8253 (MO). Amazonas:* Chachapoyas, 3—6 Km
W. of Molinopampa, 2200-2450 m, 19.vii.1962 (fl), Wurdack 1403 (F, K, NY, S,
US). *Pajonales, Pozuro, 3000 m, vii.1863 (fl), Pearce 252 (BM, K).
H. aciculare differs essentially from H. sprucei in habit and size of parts. It most
resembles some populations of the latter from Chimborazo, from which it is
separated by a disjunction of merely c. 100 km. Other, less similar populations of H.
sprucei, however, are found in the same area as H. aciculare, which occurs in two
ecologically somewhat distinct forms with widely overlapping areas of distribution. If
* Wet habitat form, see p. 68.
67
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 haud torquescentibus, petalis angustioribus, differt; a H. aciculari
Kunth isdem characteribus differt. Type: Peru, Amazonas, Chachapoyas, middle
eastern Calla-Calla slopes, near Kms 416-419 of Leimebamba-Balsas road, 2900
—3100 m, 9.vii.1962 (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 S—8 x 0-8—1-2 mm, subulate,
distally conduplicate and + cucullate, otherwise incurved with midrib slightly
prominent proximally to slightly impressed beneath, margin narrowly glaucous,
chartaceous; apex acicular, base broadened, markedly sheathing, pairs united to
form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands rather dense,
THE GENUS HYPERICUM L.
visible beneath when young. Jnflorescence 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 1-1-5mm,
narrowly-lanceolate-acerose, long-acuminate, cucullate, with golden-hyaline mar-
gin; veins 3-5, unbranched, not prominent; glands linear, interrupted in distal 1/4.
Petals golden yellow, 8-12 x 2-5-4 mm, 1-2-1-4 xX 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 (?—3300) m.
Peru (Amazonas (Chachapoyas), Junin, Pasco), apparently almost restricted to the
Cerro Calla-Calla. Map 14.
PERU. Amazonas: Chachapoyas, Cordillera Calla-Calla between Leimebamba
and Balsas, 2900-3000 m, 21.viii.1963 (fl), Ferreyra & Acleto 15284 (NY); between
Leimebamba and Balsas, 2700 m, 1.vi.1963 (fl & fr), Lépez, Sagastegui & Callantes
4429 (NY); Cerro Calla-Calla, E. side, 19 km above Leimebamba on Balsas road,
3100 m, 4.vi.1964 (fl), Hutchinson & Wright 5505 (F, NY, UC, US); Jalca de
Calla-Calla, 3100 m, 7.v.1970 (fl & fr), Sagdastegui 7451 (F, NY, VEN); Chachapoyas
to Moyobamba, La Jalca, 2700-3300 m, 20.i.1930 (fl), Williams 7583 (F, NY);
Chachapoyas, 1826 (fl & fr), Mathews 2130 (BM, K, P). Junin: Huancayo, Quebrada
Occopilla, ii.1948 (fl), Soukoup 3636 (MO). Pasco: Oxapampa, Cordillera Yana-
chaga, Cerro Pajonal 12 km SE. of Oxapampa, 2700-2800 m, 7.x.1982 (fr), Foster
8992 (BM, MO).
H. recurvum is clearly related to the southern, low, many-stemmed form of H.
sprucei, differing from it mainly in the often glandular, acuminate leaves, which
becomes recurved, not twisted, and are suffused with a golden tint (at least when
dried). The longer styles and narrower, acuminate sepals distinguish H. recurvum
from H. andinum, which has an overlapping area of distribution.
37. Hypericum wurdackii N. Robson, sp. nov.
H. recurvo N. Robson affinis, sed foliis longioribus, carinatis, densissime imbricatis,
valde tetrastichis alas formantibus, floribus maioribus, differt; a H. aciculare Kunth
isdem characteribus differt. Type: Peru, Amazonas, Chachapoyas, summit of Cerro
Malcabal (Cerro Tumbe), 3-6 km SW. of Molinopampa, 2850-2900 m, 20.vii.1962
(fl & fr), Wurdack 1426 (K!, holotype; F!, GH!, MICH!, NY!, P!, S!, US, isotypes).
Shrub 0-15—0-3 m tall, several-stemmed, erect, with branches strict, pseudo-
dichotomous and sometimes lateral. Stems orange-brown, completely enclosed by
leaves when young, eventually terete, cortex exfoliating irregularly; internodes 1
mm long. Leaves sessile, imbricate, markedly tetrastichous with the vertical rows
winglike, not or scarcely outcurving, not twisted, deciduous above the base after
turning brown; lamina 9-14 x 1-2 mm, linear-acerose, incurved-conduplicate,
cucullate, carinate wholly or only at apex and base, margin narrowly golden-hyaline
especially towards base, otherwise concolorous, yellow-green, not glaucous, char-
taceous; apex sharply acute, base broadened, markedly sheathing, pairs united to
form deep interfoliar ridge; basal vein 1, unbranched; laminar glands dense, invisible
below in mature leaf. Inflorescence 1-flowered, with pseudo-dichotomous branches
from node below; pedicel 2-3 mm long, not incrassate upwards; upper leaves not
transitional. Flowers 20-25 mm in diam., stellate. Sepals 10-12 x 3-4 mm, lan-
ceolate-acerose, 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 xX 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.
<x
70
NORMAN K. B. ROBSON
No habitat data known; 2850—2900 m.
Peru (Amazonas-Chachapoyas). Known only from type collection. Map 13 (p. 62).
PERU. Amazonas: Chachapoyas, summit of Cerro Malcabal (Cerro Tumbe), 3-6
km SW. of Molinopampa, 2850—2900 m, 20.vii.1962 (fl & fr), Wurdack 1426 (F, GH,
K, MICH, NY, P, S, US).
This bizarre species, which resembles a large Cassiope tetragona in habit, is clearly
related to the low, many-stemmed form of H. sprucei from Piura. Although it is
larger in all parts than H. recurvum, it can be interpreted as carrying the trends in leaf
form and arrangement of that species a stage further. The leaves are even more
closely imbricate, so that the tetrastichy characteristic of the young shoots of H.
recurvum is very marked in H. wurdackii.
38. Hypericum costaricense N. Robson, sp. nov.
H. sprucei N. Robson affinis, sed foliis subcoriaceis saepe glaucis, apice haud vel vix
cucullatis, floribus minoribus, sepalis apice acutissimis, stylis brevioribus, stigmati-
bus anguste vel late capitatis, capsula ovoidei-ellipsoideis, differt. Type: Costa Rica,
Alajuela, upper slopes of Volcan Pods, c. 2700 m, 1.1x.1968 (fl & fr), Wilbur & Stone
10634 (BM!, holotype; DUKE!, NY!, US!, isotypes).
Shrub or shrublet c. 0-1-1 m tall, erect or ascending, forming dense rounded clumps,
with branches strict, pseudo-dichotomous and lateral. Stems orange-brown, 4-
angled and (?) ancipitous when young, eventually terete, cortex exfoliating in strips;
internodes 2-4 mm long. Leaves sessile, + imbricate to suberect, spreading to
outcurving and twisting, + markedly tetrastichous when young, deciduous above
base without fading; lamina 5-15 x 0-6-1 mm, linear-elliptic to linear, incurved-
canaliculate, not or scarcely cucullate, midrib impressed beneath, margin narrowly
hyaline, concolorous, lucent or often glaucous, subcoriaceous; apex acute to sub-
acute, base parallel-sided, scarcely broadening, pairs forming narrow interfoliar
ridge; basal vein 1, unbranched; laminar glands + dense, visible beneath. /nflores-
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 xX 0-8-1-6 mm,
narrowly oblong to narrowly triangular-lanceolate, sharply acute, margin hyaline,
veins 5(7), unbranched, midrib or all sometimes prominent; glands linear, distally
punctiform. Petals bright yellow to orange-yellow, (6-5—)8-13 x 3-4mm, c. 1-3-1-5
X sepals, oblong-obovate; apiculus sharply acute; glands linear, distally punctiform.
Stamens c. 40-50, longest 5-6 mm long, c. 0-65 —0-75 x petals. Ovary c. 1-5-2 x
1-1-5 mm, ellipsoid; styles 3, 1-5-2 mm long, 0-7-1-5 X ovary, erect; stigmas
narrowly to broadly capitate. Capsule 4-5 x 2-5-3 mm, ovoid-ellipsoid, obtuse to
rounded, shorter than sepals. Seeds 0-7-1 mm long, ecarinate; testa finely scalar-
iform-reticulate.
In open paramo on rocky, often volcanic slopes; (900—)2440-3700 m.
Costa Rica (San José, 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, 1.ix.1968 (fl &
fr), Wilbur & Stone 10634 (BM, DUKE, GH, MO, NY, US). Cartago: Asuncién of
Cerro de la Muerte, c. 3150 m, 17.v.1971 (fl), Wilbur 14563 (DUKE, MICH, MO).
Limon: Chirrip6 National Park, between Casa da Administraci6n 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 et al. 2389 (BM, MO).
PANAMA. Bocas del Toro: Valle de Silencia, 2440 m, 8.viii.1979 (fl & fr),
Antonio 1547 (BM, MO). Chiriqui: Cerro Fabrega and vicinity, near Costa Rican
frontier, 3150—3335 m, 7—8.iv.1976 (fl), Weston 10149 (MO).
COLOMBIA. Antioquia: Paramo Frontino, near Llano Grande, 3450 m,
25.x.1976 (fl & fr), Boeke & McElroy 229 (NY), Bolivar/Antioquia: Paramo de
Chaquiro, 900—960 m, 23.ii.1919 (fl & fr), Pennell 4285 (GH, NY).
THE GENUS HYPERICUM L.
71
Map 15 _ 38. H. costaricense @; 39. H. bryoides O; 44. H. marahuacanum: a. subsp. marahuacanum @, b.
subsp. strictissimum A, c. subsp. chimantaicum OQ.
H. costaricense is related to 34 H. sprucei, having similar markedly incurved leaves
that twist when long enough to do so, but they are not cucullate and no more than
subcoriaceous. Moreover, the flowers are smaller, with shorter styles and capitate
stigmas.
In a specimen from the highest point of the Pan-American Highway on the
Talamanca Range (J. & C. Taylor 11757 (NY)) the leaves are broader, the flowers
larger, and the styles longer than in the other specimens. It seems probable that the
suggestion on the label is correct, i.e. that it is the hybrid H. irazuense x H.
costaricense. The two Colombian collections belong to H. costaricense despite the
disjunct distribution and the lower altitude of Pennell 4205, which is a dwarf shrub
(0-1—0-15 m tall) with numerous lateral branches, leaves relatively long (c. 10 mm),
recurving markedly and twisting, flowers that are relatively small for the species but
nevertheless fall within its range of variation, and stigmas that are broadly capitate. It
resembles a dwarf version of the (primitive) Panamanian form of H. 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, Santandér,
Paéramo de Santurban, 3000 m, 18.11.1927 (fl), Killip & Smith 19533 (NY!,
holotype; GH!, US!, isotypes).
Suffrutex 0-09-0-15 m tall, erect or ascending from creeping branching and rooting
base, forming dense Polytrichum-like rounded clumps, with branches strict, pseudo-
dichotomous and lateral. Stems orange-brown, 4-angled and(?) ancipitous when
young, soon terete, cortex exfoliating irregularly; internodes 1-3-5 mm long. Leaves
sessile, imbricate, eventually outcurving and slightly twisting, tetrastichous, de-
ciduous above base without fading; lamina 3-5 x 0-—0-8 mm, linear-subulate,
incurved-canaliculate, cucullate, midrib impressed beneath or not, margin narrowly
hyaline, concolorous, not glaucous, chartaceous; apex subacute to obtuse, base
gradually broadened and sheathing, pairs forming relatively broad interfoliar ridge;
basal vein 1, unbranched; laminar glands dense, visible beneath. Inflorescence
1-flowered, terminal, with pseudo-dichotomous branches from node below; pedicel
c. 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
,
ip
NORMAN K. B. ROBSON
oblong-lanceolate, acute, incurved above, margin hyaline, veins 3—5, unbranched,
midrib or all prominent; glands all linear or distally punctiform. Petals bright yellow,
3-5-6:5 X 1-5-3 mm, c. 1:2—1-4 X sepals, obovate; apiculus acute or obsolete;
glands striiform. Stamens 20-35, longest 2-5-3 mm long, c. 0-75 x petals. Ovary
0-8-1 x 0-6—0-8 mm, ellipsoid; styles 3, 1-1-2 mm long, c. 1-2 X ovary, erect;
stigmas narrowly to broadly capitate. Capsule 4-4-5 x 2-2-5 mm, cylindric-
ellipsoid, subacute to obtuse, slightly exceeding sepals. Seeds not seen.
Rocky places in open paramo; 3000-4200 m.
Colombia (Norte de Santandér, Santandér, Boyacd). Map 15 (p. 71).
COLOMBIA. Boyaca: Sierra Nevada del Cocuy, Alto La Cueva, 3850 m,
5.vi.1973, Cleef 10000 (BM, COL, U). Norte de Santandér: Paramo de Romeral,
3800-4200 m, 30.i.1927 (st), Killip & Smith 18631 (F, GH, K, NY, S, US); Péaramo
de Santurban, 5 km E. of Berlin, 3200 m, 4.vii.1984 (fl), Wood 4504 (K). Santandér:
Paéramo de Almorzadero, 3500-3700 m, 20.vii.1940 (fl), Cuatrecasas & Garcia
Barriga 9993 (F), 9995 (COL), 9996 (NY).
H. bryoides is a reduced suffruticose version of the more specialized form of H.
costaricense, differing from it essentially in size of parts. The dense moss-like clumps
expand by the creeping and rooting of the outer shoots.
40. Hypericum bolivaricum N. Robson, sp. nov.
H. magdalenico N. Robson affinis, sed foliis angustissime ellipticis basi angustatis
saepe terminantibus, petalis brevioribus, staminibus paucioribus, stylis brevioribus
stigmatibus manifeste capitatis, differt. Type: Colombia, Bolivar, headwaters of Rio
Sinu, 1918 (fl), 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 3 x 2 mm,
narrowly ellipsoid; styles 3, 4-5 mm long, 1:5 X ovary, slightly outcurved; stigmas
rather broadly capitate. Capsules and seeds not seen.
No habitat or altitude cited.
Colombia (Bolivar). Known from only one collection. Map 16 (p. 74).
COLOMBIA. Bolivar: headwaters of Rio Sinu, 1918 (fl), Pennell 4783 (NY).
Although only one specimen of H. bolivaricum has so far been studied, its taxonomic
position is clear and it is quite distinct from its nearest relatives. Morphologically and
geographically it is intermediate between the long-leaved form of H. magdalenicum
and the H. juniperinum group, having the thickly coriaceous leaves of the former but
the 1-nerved, twisting leaves and predominantly lateral flowers of the latter.
41. Hypericum juniperinum Kunth
in Humboldt, Bonpland & Kunth, Nova Gen. et Sp. Pl. 5: 189 (1822); Bentham, Pl.
Hartweg.: 165 (1845). Type: Colombia, Cundinamarca ?, ‘in frigidis Andium
Nova-Granatensium’, vii.1805 (fl & fr), Humboldt & Bonpland (P-HUM!, holo-
type; P! (F!, photograph), GH!, isotypes).
THE GENUS HYPERICUM L.
Brathys juniperina L. f., Suppl. Pl.: 268 (1781). Type: Colombia, Cundinamarca, ‘in
Nova Granada’, Mutis (LINN!, holotype; BM!, MA!, US!, isotypes).
H. brathys Sm., Plant. Ic. Ined. Herb. Linn. 2: 41, t. 41. (1790), nom. illegit.; Lam.,
Encycl. méth. (Bot.) 4: 152 (1797); Choisy, Prodr. monogr. fam. Hypéric.: 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 Annis Sci. nat. (Bot.) IV, 18: 292
(1862); R. Keller in Bull. Herb. Boissier II, 8: 177 (1908), in Engler & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 181 (1925); Knuth in Reprium Spec. nov. Regni veg.
Beih. 43: 484 (1927); Gleason in Bull. Torrey 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.] B 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. vég. 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, Mérida,
‘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 Batallén, 3000 m, iii.1911 (fl), Jahn 2 (B+, 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 787 (NY!, holotype; GH!, K!, isotypes).
H. strictum var. reductum Gleason in Bull. Torrey bot. Club 56: 104 (1929). Type:
Colombia, Santandér, west slope of the Paramo de Santurban towards Tona, 3000
m, 18.ii.1927 (fl & fr), Killip & Smith 19534 (NY!, holotype).
Shrub or shrublet 0-2—2-5 m tall, erect or rarely decumbent and rooting at the base,
with branches strict, lateral or sometimes pseudo-dichotomous. Stems orange-
brown, 4-angled and ancipitous when young, soon 2-lined, eventually terete, cortex
exfoliating irregularly; internodes 1-5—10 mm long. Leaves sessile, not or scarcely
imbricate, erect to spreading and twisting, not or scarcely tetrastichous, deciduous
above base without fading; lamina 6-14 x 0--1-5 mm, linear to acicular, rarely
cucullate, midrib impressed beneath, margin narrowly hyaline, concolorous, rarely
glaucous, coriaceous to subcoriaceous; apex acute, base parallel-sided, scarcely
broadening, pairs forming narrow interfoliar ridge; basal vein 1, unbranched;
laminar glands dense to sparse, sometimes visible beneath. Inflorescence 1-flowered,
terminal (usually) and on up to 12 short lateral shoots, sometimes with pseudo-
dichotomous branches from terminal node; pedicel 0-5-3 mm long, not incrassate
upwards; upper leaves transitional. Flowers 4-12 mm in diam., stellate to obconic.
Sepals 2-5-7 x 0-5-2 mm, narrowly lanceolate, acute, veins 5, unbranched, midrib
not prominent, margin hyaline; glands linear, sometimes distally punctiform. Petals
bright to pale yellow, (3-)4-9 x 1-5—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 Xx ovary, ascending; stigmas narrowly to broadly capitate.
Capsule 2:5—4 x 2-3 mm, ovoid-subglobose to globose, shorter than sepals. Seeds
0-6—1-2 mm long, ecarinate; testa finely scalariform.
73
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 subpéramo;
(1800) 2200-3800 m.
Venezuela (Trujillo, Mérida, Tachira), Colombia (Magdalena and Norte de Santan-
dér 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.1938 (fl & fr), Cuatrecasas
1197 (F); Sierra Nevada del Cocuy, between Praton and Cobugon, c. 3100 m,
15.viii.1957 (fl), Grubb, Curry & Fernandez-Perez 507 (COL, K); Guican, Hoya del
rio Tabor, en direccione a Ritacuva, 3570-3530 m, 18.ix.1969 (fl), Cuatrecasas &
Rodriguez 27833 (BM, COL, US); Paramo de la Rusia, NW.-N. de Duitama, cerca
del puente de rio Surba, 3240 m, 9.xii.1972 (fl & fr), Cleef 6925 (COL, U); carreteria
Sogamoso-Pajarito, Km. 57-73, 2850-3010 m, 20.ix.1969 (fl), Diaz 196 (COL).
Caldas: cabeceras del rio Ottin, 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 Retiro,
Bogota, 2600-2700 m, (v.1946 (fl & fr), Schultes 7004 a (COL, F, GH); Bogota to
Choachi, 27 km W. of Choachi, 3260 m, 4.i.1976 (fl), Luteyn 4730 (BM, COL, MO,
NY); Paéramo de Guasca, 2840 m, 15.xii.1938 (fl), Balls 5680 (BM, COL, F, K);
Paramo de Sibaté, 3570 m, 22.xi.1981 (fr), Inglesias 19 (COL); entre Bogota y La
Calera, 2650-3000 m, 27.xi.1947 (fl), Barkley, Garcia-Barriga & Vanegas 17C814
(COL, US). Magdalena: Sierra Perija, near Laguna el Juncos, above ‘Africa’, c.
2200 m, 16.xii.1944 (fl & fr), Haught 4503 (F, K). Meta: Paramo de Sumapaz, Hoya
El Nevado, Laguna La Guitarra, 3460 m, 21.i.1972 (fr), Cleef 824 (COL, U). Norte
de Santandér: Péramo 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 Péramo de Taébano, c. 2800-3000 m, 1.1.1946 (fl), Schultes & Villareal
7834a (F). Santandér:* Bucaramanga (Cumbre to Cicuta), 3300 m, v.1948 (fl),
* Robust, large-flowered form (= H. pseudobrathys Turcz. sensu stricto).
THE GENUS HYPERICUM L.
Sandeman 6067 (BM, K); Paramo de Almorzadero, 3600-3800 m, 28.xi.1941 (fl),
Cuatrecasas 13493 (COL, F, NY); limites entre los Departamentos Santandér 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. Mérida: Péramo de La Negra above Bailadores, 3100 m,
13.ii.1939 (fl), Alston 7024 (BM, S, U);* Paramo de Muruti, x.1865 (fl), Moritz 1165
(BM, GH, K, P, S); Paramo La Negra between Tovar and La Grita, 3000 m, 7.x.1965
(fl), Breteler 4619 (K, MO, NY, S, U, VEN). Tachira: Paramo Zumbador, 14km S.
of El Cobre, 2500 m, 31.iii.1974 (fl), Gentry, Morillo & Morillo 11068 (GH, MO,
NY, VEN); Dtto. Uribante, a las cabeceras del rio Uribante, 51 km al SW. de
Pregonero, 2700-2900 m, 28.ix.1981 (fl), Steyermark & Manara 125420 (BM, VEN).
Trujillo: Alredores de Guirigay, hacia Laguna La Parida, 3300 m, viii.1958 (fl),
Aristeguiéta & 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,
Mérida) and in isolated Colombian localities in Santandér, Boyaca, Cundinamarca,
Antioquia, and Cauca, but it is linked to the typical ‘H. brathys’ by intermediates.
Indeed Kunth’s type of H. juniperinum has flowers somewhat intermediate in size.
In Cauca there is a separate apparently continuous morphocline in which the styles
are always three in number. This cline links the tall erect plants with crowded
subappressed leaves and larger flowers (i.e. H. juniperinum sensu Spach and Triana
& Planchon) with smaller, spreading to decumbent plants with stems rooting, longer
internodes, smaller, closely appressed leaves, and smaller flowers. In its extreme
expression, the latter form approaches H. prostratum, but can be distinguished from
it by the larger leaves and flowers and the longer styles. Specimens exemplifying this
end of the morphocline include Idrobo, Pinto & 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.1952 (fl), Kgie 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(—1-2) mm,
linear to acicular, cucullate, midrib impressed beneath, margin broadly hyaline or
golden, concolorous, not glaucous, subcoriaceous, apex acute, base parallel to
cuneate, not broadening, pairs forming narrow interfoliar ridge; basal vein 1,
unbranched; laminar glands + dense and prominent above, sparse or not visible
beneath. Inflorescence 1-flowered, terminal and sometimes on up to 6 short lateral
shoots, without pseudo-dichotomous branches; pedicel almost absent or up to 2 mm
long, not incrassate upwards; upper leaves not transitional. Flowers 4-8 ,m in diam.,
stellate to obconic. Sepals 4-5, 2-3-5 X 0-5-1 mm, lanceolate, acute, veins 5,
unbranched, midrib not prominent; glands linear. Petals 4-5, bright yellow, 3-6 x
1-5-2 mm, 1-5-2 xX 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 0-8mm,
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.
ip)
76
NORMAN K. B. ROBSON
Colombia (Cundinamarca, Meta, Boyaca, Santandér). 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.1972 (fl
& fr), Cleef 6747 (COL, U); between Soata and Cocuy, Paramo del Alto del Escobal,
3750 m, 8.ix.1938 (fl), Cuatrecasas 1224 (F). Cundinamarca: Bogotaé-Usmé road,
Paramo de Chisaca, between Kms 28 and 33, 3375-3450 m, 17.v.1979 (fl), Luteyn
7770 (BM, COL, MO, NY); Paéramo de Cruz Verde, aprox. 5 km al ENE. de la
Laguna El Verjoén, 3360 m, 2.v.1972 (fl), Cleef 3371 (BM, COL, U); entre Cogua y
San Cayetano, Laguna Verde y alredores, 3600 m, 12.xi.1972 (fl), Cleef 6260 (BM,
COL, U). Meta: Paramo de Sumapaz, Hoya El Nevado, Laguna La Guitarra y
alredores, 3460 m, 21.1.1972 (fl), Cleef 834 (BM, COL, U). Santandér: Péaramo de
Almorzadero, 3500-3700 m, 20.vii.1940 (fl), Cuatrecasas & Garcia Barriga 9998
(COL, US).
H. prostratum continues one of the reduction trends in H. juniperinum, differing
from the decumbent Cauca plants in (for example) its prostrate habit, shorter and
initially strongly appressed leaves, and smaller flowers and fruits. The direction of
the trend, however, has been reversed, i.e. the westernmost specimens (from
Cundinamarca and Meta) are nearest to the above-mentioned Cauca populations of
H. juniperinum both morphologically and geographically.
43. Hypericum parallelum N. Robson, sp. nov.
H. magdalenico N. Robson affinis, sed foliis angustioribus glandulis prominentibus
costa media haud ramosa, floribus minoribus stylis brevioribus sepalis glandulose-
punctatis, differt; a H. pimelioidei Planchon et Linden ex Triana et Planchon inter
alia ramis strictissimis foliis angustioribus valde tetrastichis differt. Type: Colombia,
Norte de Santandér/César, Linea divisoria entre los Deptos. Santandér del Norte y
César, 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 Santandér/César); known only from Las Jurisdicciones, Cerro de
Oroque. Map 17.
COLOMBIA. N. de Santandér/César: 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.
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 haud ramosa, sepalis angustioribus stylis gracilioribus brevioribus, inter
alia differt; a H. lancifolio Gleason foliis angustioribus lucentibus torquescentibus
margine valde incurvatis differt. Type: Venezuela, Amazonas, Dept. Atabapo,
Cerro Marahuaca, cumbre Seccion suroriental, 2685 m, 15.i.1981, Maguire, Steyer-
mark et al. 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 exfoliating irregularly; inter-
nodes 0-7—2 mm long. Leaves sessile, densely imbricate, erect to suberect, outcurv-
ing, twisting slightly or not, subtetrastichous to strictly tetrastichous, deciduous
above base without fading; lamina 5—12 x 0-6-1 mm, linear, incurved, not cucullate,
midrib impressed beneath, margin narrowly hyaline, concolorous, polished, not
glaucous, coriaceous; apex acute, often slightly pungent, base parallel-sided, not
sheathing, pairs united to form narrow interfoliar ridge; basal vein 1, without
reticulation; laminar glands in single irregular row, visible beneath or not. Jnflores-
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 2-5-4 mm, c. 1-2—1-7 X sepals, obovate (?) to oblanceolate; apiculus
sharply acute; glands linear, usually distally interrupted. Stamens 30-50, longest 3-9
mm long, 0-45—0-8 x petals. Ovary 1-2-3 x 1-1-5 mm, narrowly ellipsoid or
narrowly ovoid-ellipsoid to ellipsoid-subglobose, acute to rostrate; styles 3(4), 2-5
mm long, 1-5-2 X ovary, outcurved to erect or ascending; stigmas narrow to
narrowly capitate. Capsule 3-5-5 x 2-2-5 mm, narrowly ovoid to ovoid-ellipsoid,
shorter than sepals. Seeds 1-1-2 mm long, ecarinate; testa finely scalariform.
Along shaded (?) streams, in swampy areas, and in drier places and on escarpment;
2500-2800 m.
Colombia (Boyacé), Venezuela (Mérida, Amazonas, Bolivar). Map 15 (p. 71).
|
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78 NORMAN K. B. ROBSON
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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 %2; b-g X 3). A. Maguire et al. 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 (Mérida, Boyaca) the inflorescence branching is
wholly lateral.
THE GENUS HYPERICUM L.
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 < 1 mm, with glands visible
beneath. /nflorescence branches pseudo-dichotomous; pedicel 2—3 mm long. Flow-
ers 10—15(?-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 2mm,
narrowly ovoid.
Along shaded (?) steams and on escarpment; 25—2800 m.
Venezuela (Amazonas).
VENEZUELA. Amazonas: Departamento Atabapo, Cerro de Marahuaca, parte
central de la Meseta S.-E., Quebrada Yekuana, afluente del rio Negro, 2560 m,
10—12.x.1983, Steyermark 129511 (BM, VEN); Cerro de Marahuaca, al NE. de, y
casi contigua con, Cerro Duida, c. 2750 m, 2 & 9.ii.1975, Tillett, Colveé et al. 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.1944 (fl), Fosberg 22209 (US!, holotype; NY!, P!,
isotypes).
Icon: Fig. 12B.
Branches strict to very strict, some paired and pseudo-dichotomous or all lateral.
Leaves twisting slightly or not, markedly tetrastichous; lamina 5—11 x 0-5—0-8 mm,
with glands relatively few, mostly obscure. Inflorescence branches short, lateral,
from up to c. 6 nodes below, sometimes with pseudo-dichotomous branches from
uppermost node; pedicel c. 2 mm long. Flowers c. 10-12 mm in diam. Sepals 4—6 x
0-7—1-5 mm, narrowly lanceolate, veins (3)5. Petals 7-9 x 3-4 mm, c. 1-7 X sepals,
obovate (?). Stamens c. 40, longest c. 3-4 mm long, c. 0-45 x petals. Ovary 1:2-1-5
x 1 mm, ellipsoid-subglobose; styles 2-5-3 mm long, c. 2 X ovary, suberect; stigmas
scarcely enlarged. Capsule 4 x 2:5 mm, ovoid, acute, shorter than sepals.
Paramo and subparamo, in dry stony places; 2700-3250 m.
Colombia (Boyaca), Venezuela (Mérida).
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 Belén, c. 14kmN. of Belén, near 259 km marker (from Bogota), c. 3250
m, 9.v.1959 (fl), Barclay & Juajibioy 7668 (NY).
VENEZUELA. Mérida: Péramo de Las Coloradas (entre Santa Cruz de Mora y
El Molino), San Rafael, 2950-2950 m, 9.ii.1973 (fl), Cuatrecasas, Ruiz-Teran &
Lépez-Figueiras 28529 (BM, US); Sierra Nevada, i.1865 (fl), Moritz 1602 (BM).
Subsp. strictissimum, which comprises two geographically distinct populations,
differs from subsp. marahuacanum in having narrower leaves, smaller flowers which
terminate lateral branches as well as main ones, and shorter styles. When well
719
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
haud vel vix punctata, pedicellis longioribus, floribus minoribus interdum ramulis
brevibus lateralibus terminantibus, stylis brevioribus anguste capitatis, capsula
elliptica, differt. Type: Venezuela, Bolivar, Chimanta Massif, Central Seccion, E.
branch of headwaters of Rio Tirica, 2121 m, 12.11.1955 (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. /nflorescence sometimes with 2—4 short axillary shoots
immediate below, pseudo-dichotomous; pedicel c. 3 mm long. Flowers c. 10 mm in
diam. Sepals 4-6 x 0-6-1 mm, linear-lanceolate, acute, veins 5-7. Petals 6-8 xX
2-5-3 mm, c. 1:5 X sepals, oblanceolate. Stamens c. 30?, longest c. 4—-S 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., Apacarda-tepui,
sector Norte de Macizo, c. 2200 m, 30.i—1.ii.1983 (fl), Steyermark, Huber & Carreno
128351 (BM, VEN), Huber & Steyermark 7014 (BM, VEN); Cimo del Roraima,
i.1977 (fl), Delascio & Brewer 4794 (VEN).
All the characters by which subsp. chimantaicum differs from subsp. marahuacanum
are apomorphic (advanced). It is the easternmost taxon of those in the H.
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, Santandér,
vicinity of La Baja, 2200-2600 m, 14.i.1928 (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.
from node below; pedicel 2—4 mm long, rather slender; upper leaves not transitional.
Flowers 10-20 mm in diam., stellate. Sepals 6-10 x 1-5-3 mm, narrowly to
acuminate, incurved above, veins 3—5, unbranched or laterals branched, midrib not
prominent; glands sometimes linear to striiform towards base, otherwise (or wholly)
punctiform. Petals pale yellow, 8-12 x 3-5 mm, 1-1-1-3 X sepals, narrowly
obovate; apiculus acute; glands punctiform. Stamens c. 30-70, longest 5-7 mm long,
0-5—0-75 X petals. Ovary 2:3 X 1-2 mm, ellipsoid; styles 3, 2-4 m long, 1-1-75 x
ovary, Outcurving; stigmas scarcely to broadly capitate. Capsule 4—7 x 2-5-3 mm,
ellipsoid, acute, shorter than sepals. Seeds 0-7—0-8 mm long, ecarinate; testa finely
scalariform-reticulate.
In paramo; c. 2600-3700 m.
Colombia (Boyacé, Santandér, N. de Santandér), Venezuela (Tachira, Apuré).
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 Santandér:
Pamplona, cerro al NE., 2700 m, 26.vii.1940, Cuatrecasas & Garcia Barriga 10218
(COL, F, US). Santandér: Paramo de la Puentas, above La Baja, 3500-3700 m,
25.i.1927, Killip & Smith 18187 (F, GH, K, NY, S, US).
VENEZUELA. Apuré: Paéramo 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.i.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 H. magdalenicum, differing
in the more slender stems, the narrower smaller, thinner leaves, and the smaller
flowers with shorter styles. The typically plane narrow leaves are distinctive.
H. lancifolium has usually been labelled H. weberbaueri, but it differs from that
Peruvian species (= H. struthiolifolium) in several respects, e.g. the plane (not
incurved) leaves with prominent glands. These two species are not closely related.
The Venezuelan population has shorter, narrower, outcurving leaves and smaller
flowers with fewer stamens and shorter ovary, styles, and capsule. In addition, its
height is usually less and its sepals usually narrower. The leaves are rarely plane as in
H. lancifolium, always more or less incurved. It thus forms a transition to 46 H.
horizontale.
46. Hypericum horizontale N. Robson, sp. nov.
H. lancifolio Gleason affinis, sed habitu prostato vel adscendenti, foliis secundis
apice acicularibus, floribus minoribus, stigmatibus vix vel anguste capitatis, capsula
minori, inter alia differt. Type: Colombia, Santandér, Paramo de Las Vegas,
3700-3800 m, 20—21.xii.1926, Killip & Smith 15617 (BM!, holotype; A!, GH!, K!,
US!, isotypes).
Shrublet, up to c. 0-05 m tall, matted, with branches up to c. 0-3 m long, prostrate
with apex ascending, rooting, branching mainly pseudo-dichotomous. Stems orange-
brown, 4-angled and ancipitous when young, soon 2-lined, eventually terete, cortex
exfoliating irregularly; internodes 1-3 mm long. Leaves sessile, not or scarcely
imbricate, ascending to outcurving and secund, not tetrastichous, deciduous above
base without fading; lamina 5-9 x 0-5—1-3 mm, linear, incurved-canaliculate, not
cucullate, midrib impressed dorsally, margin narrowly hyaline, concolorous when
mature, often red-tipped when young, sometimes sublucent, not glaucous, sub-
coriaceous; apex acicular, base parallel-sided, scarcely broadened, pairs forming
narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense to sparse,
visible on both sides. Inflorescence 1-flowered, terminal, and sometimes also on up to
9 short, paired lateral shoots, usually with pseudo-dichotomous branches from
terminal node; pedical 1-2 mm long or almost absent; upper leaves not transitional.
Flowers 8-12 mm in diam., stellate (? or shallowly obconic). Sepals (3-5—)4-6
— X 1-1-5 mm, narrowly oblong to narrowly or broadly lanceolate, distally incurved,
82
NORMAN K. B. ROBSON
acute, margin hyaline, tip sometimes reddish, veins 5, unbranched, prominent;
glands linear towards base, punctiform or absent distally. Petals bright (?) yellow,
5-7 X 2:5-3-5 mm, 1-1-1-4 X sepals, obovate; apiculus apiculate; 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 Santandér, Santandér). Map 18.
COLOMBIA. Norte de Santandér: Paramo de Santurban, en route from Tona to
Mutiscua, 4200-4500 m, 19.ii.1927 (fl & fr), Killip & Smith 19564 (BM, GH).
Santandér: Paramo de Las Vetas, a 2 km del paraje de Berlin, c. 2800 m, 18.xii.1948
(fl & fr), Molina & Barkley 1885410 (COL, F); idem, c. 3450 m, 3.vi.1960, Barclay &
Juajibioy 10445 (MO, NY); Paramo de Almorzadero, 4200 m, 9.ix.1955 (fl),
Vareschi 4070 (VEN p.p.); idem, auf der Passhéhe, 3850 m, 24.vii.1952, Schneider
1232 (Si);
Although quite different in habit, H. horizontale is clearly related to the Venezuelan
population of H. lancifolium. Its distinct distribution and morphological differences,
taken together, indicate that it should be treated as a separate species.
47. Hypericum tetrastichum Cuatrec.
in Ciencia Mex. 23: 144, f. 3A—G (1964). Type: Colombia, Santandér, Municipio de
Onzaga, vereda de Chaguaca, alto de la Laguna de Los Bobos, en el filo divisorio
Santandér-Boyaca, 3800 m, 7.viii.1958 (fl), Jaramillo-Mejia, Herndndez-
Camacho & van der Hammen 922 (US, holotype; COL, isotype); same data,
Jaramillo-Mejia, Hernaéndez-Camacho & van der Hammen 918 (COL!, topotype).
Shrub or shrublet0-05—1 m tall, erect and + densely caespitose, with branches strict,
pseudo-dichotomous or (mostly) lateral. Stems orange-brown to blackish, 4-lined
and compressed when young, the subfoliar ridges broad, soon or eventually terete,
cortex exfoliating in strips or irregularly; internodes 0-5-6 mm long. Leaves sessile,
erect, densely imbricate, markedly tetrastichous, the longer eventually spreading or
Map 18 46. H. horizontale A; 47. H. tetrastichum @.
THE GENUS 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, cylindric-ellipsoid to ellipsoid-subglobose, obtuse to round-
ed, shorter than sepals. Seeds 0-6—0-8 mm long, ecarinate; testa finely scalariform.
Wet or dry, often exposed areas in paramo and subparamo; 2800-4160 m.
Colombia (Boyaca, Santandér, Venezuela (Mérida). Map 18.
COLOMBIA. Boyaca: Municipio Villa de Leyva, Parque Nacional Santuario de
Iguaque, 3700 m, 20.vii.1979, Melampy 161 (COL); Municipio de Duitama, Paramo
de La Rusia, 3400-3500 m, 28.xi.1978 (fl), Diaz P. 1243 (COL); Paramo de
Guantiva, 3100 m, 26.vi.1964, Espinal 1733 (COL); paramos al NW. de Belén,
vereda S. José de la Montana, Alto de las Cruces y alredores, Cabeceras Q. El Toral,
c. 3950 m, 24.ii.1972 (fl), Cleef 1736 (COL, U). Santandér: Paramo Ruso, 3150 m,
12.vii.1968 (fl), Barclay & Mullen 38C137 (COL).
VENEZUELA. Merida: Sierra Nevada de Mérida, margen N. de la Laguna verde
(al pie del Pico Humboldt), + 4100 m, fl. 15-18.ii1.1966, Schulz, Rodriguez &
Ramirez 358 (VU).
H. tetrastichum is closely related to H. lancifolium and H. strictum, differing
essentially from the latter in its markedly tetrastichous foliage and in the length of the
styles. The leaves appear pungent but may not be rigid when alive.
In some Boyaca localities, H. tetrastichum occurs in two apparently distinct forms:
(i) leaves and flowers larger; leaves glandless beneath, apex acuminate-acicular
(damp habitats); (ii) leaves and flowers smaller; leaves glandular beneath, apex
acute (dry habitats). Until more information is available on the nature of this
variation (genetical or environmental), it seems preferable not to give form (ii) any
taxonomic recognition. Specimens of this form include:
Paramo de La Rusia, NW.-N. de Duitama, Laguna Negra, 3725 m, 14.xii.1972
(fl), Cleef 7208 (COL, U), 7208A (COL); Paramos al NW. de Belén, cabeceras
Quebrada Minas, Hoya Laguna El Alcohol, 3900 m, 26.11.1972 (fl), Cleef 1844 (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.
48. Hypericum strictum Kunth
in Humboldt, Bonpland & Kunth, Nova Gen. et Sp. Pl. 5: 190 (1822); Gleason in
Bull. Torrey bot. Club 56: 104 (1929) pro parte excl. vars. Type: Columbia,
Cundinamarca, prope Santa Fé 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
U
84
NORMAN K. B. ROBSON
lateral, sometimes decumbent and rooting at the base. Stems orange-brown, 4-
angled and ancipitous when young, then 2-lined, subquadrangular, eventually
terete, cortex exfoliating in strips; internodes 1-7 mm long. Leaves sessile, imbri-
cate, outcurving or spreading but not twisting, not or obscurely tetrastichous,
deciduous above base without fading; lamina 5-13 x 5-6—1-3 mm, linear-acicular,
incurved-canaliculate, cucullate or not, midrib impressed dorsally, margin narrowly
hyaline, sometimes red-tipped when young, otherwise concolorous, dull to sub-
lucent on both sides, glaucous when young, coriaceous to chartaceous; apex acute to
subacuminate, base parallel-sided, pairs forming narrow interfoliar ridge; basal vein
1, unbranched; laminar glands absent or rarely sparse and visible beneath. /nflores-
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 (Santandér, 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 H. tetrastichum, and the sepal form and glands are distinct. H.
strictum also has a different, though overlapping distribution, being centred in
Cundinamarca and showing trends north-east and south-west from there along the
Cordillera Oriental. Some Boyaca specimens verge toward H. tetrastichum morpho-
logically; others are quite distinct from that species and, indeed, form a derivative
subspecies of H. strictum.
48a. H. strictum subsp. strictum
H. struthiolifolium var. strictum (Kunth) Choisy in DC., Prodr. syst. nat. regni veg. 1:
553 (1824) [‘struthiolaefolium’|; Weddell, Chloris Andina 2: 271 (1857); Triana &
Planchon in Annis 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 Cémbita, 3445 m, 1.ix.1944 (fl),
St. John 20695 (NY, US); NW. of Belén, 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 (fl), Barclay 4157 (COL); Zipaquira — Pacho, Paramo
Alto, 11.11.1951 (fl), Romero-Castaneda 2282 (COL, F); Paramo de La Calera,
3000-3300 m, 25.ii.1950 (fl & fr), Philipson, Idrobo & Fernandez 2440 (BM, COL,
F); Bogota, paramo above El Chicé, 3150-3300 m, 11.vii.1943 (fl), Fosberg &
THE GENUS HYPERICUM L.
85
Map 19 48. H. strictum: a. subsp. strictum Ml, 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.1972 (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, Santandér, 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.] € gracile Triana & Planchon [in Weddell, Chloris Andina 2:
271 (1857), nomen] in Annls Sci. nat. (Bot.) IV, 18: 294. (1862), [‘struthiolae-
folium’| nom. illegit., pro parte, quoad spec. Linden 1313.
H. laricifolium var. glaucum R. Keller in Bull. Herb. Boissier II, 8: 182 (1908). Type:
Colombia, Boyaca, prov. Tunja, Paramo de Coati, prés 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. Param. Colomb. Cord. Or.: 304 (1981),
nomen.
Stems usually branched from the base, + ascending. Leaves S—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 1-1-2(-1-5) 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
U
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, Santandér).
COLOMBIA. Boyaca: Péramo de Santa Rosa, entre Santa Rosa de Viterbo y
Cerinza, 2800 m, 17.vii.1940 (fr), Cuatrecasas & Garcia Barriga. 9721 (COL, K, NY,
P); Paramo de Guantiva, Alto de Canutos, 3200-3400 m, 3.x.1940 (fl & fr),
Cuatrecasas 10329 (COL, F, US). Paéramo de La Rusia, NW.-N. de Duitama,
Aislada, 3490 m, 8.xii.1972 (fl & fr), Cleef 6835 (U). Santandér: Paramo de
Santurban, near Vetas, 3950—4160 m, 17.i.1927 (fl), Killip & Smith 17567 (A, BM,
K, US).
Subsp. compactum differs from subsp. strictum in the subacute to acute, cucullate,
usually glandular leaves and the markedly ribbed sepals, both leaves and sepals being
frequently red-tipped or red-tinged when young. It comprises two forms, which are
geographically almost distinct and differ mainly in habit. The form in Santandér
(‘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. Santandér del
Norte y César, 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—S(-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 xX 2 mm,
cylindric-ellipsoid; styles 3, 6-7 mm long, c. 1-5 X ovary, distally outcurved; stigmas
scarcely capitate. Capsule c. 8 X 4-5 mm, cylindric-ellipsoid to cylindric, shorter
than sepals. Seeds not seen.
THE GENUS HYPERICUM L.
Paramo; 2600-3960 m.
Costa Rica (Lim6n), Colombia (Magdalena, César/Norte de Santandér). Map 20.
COSTA RICA. Lim6n: 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.1977 (fl), Jaramillo et al. 5371 (BM, COL); loc. cit., 3300 m,
9. viii. 1977 (fl), Jaramillo et al. 5454 (COL, U). César/Norte de Santandér: 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.
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,
87
Map 20 49. H. jaramilloi A; 50. H. cardonae @;51. H. caracasanum: a. subsp. caracasanum @, b. subsp.
turumiquirense UO).
88
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 (1—)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. Jnflorescence 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 2-4mm,
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 José), Venezuela (Trujillo, Mérida, Tachira, Apuré),
Colombia (Norte de Santandér, Santandér, Boyacé, Arauca). Map 20 (p. 87).
COSTA RICA. Cartago: Cordillera de Talamanca, ‘Cerro de la Muerte’, Pan-
American Highway 5 km above Millsville (8 km above Nivel), 3400-3300 m, vii.1949
(fl), Holm & Iltis 454 (F, GH, K, MO, NY, P, U). San José: Valle de los Conejos
(upper Rio Talari) and trails to Cerro Cirrip6 and Valle de los Lagos, 3400-3820 m,
22.viii.1971 (fl), Burger 8281 (BM, DUKE, F).
COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada E] Playén, 3350 m,
13.iii.1973 (fl), Cleef 9182D (COL). Boyaca: paramos al NW. de Belén, cabeceras
Quebrada Minas, Hoya Laguna El Alcohol, 3800 m, 26.ii.1972 (fl), Cleef 1870 (U);
Cocuy Mts, valley of Quebrada Playitas above Guican, 3800 m, 24.vi.1984 (fl),
Wood 4453 (K). Norte de Santandér: Paramo de Tama, alredores de la Cueva, 3000-
3200 m, 28.x.1941 (fl), Cuatrecasas, Schultes & Smith 12691 (COL). Santandér:
Paramo de Santurban near Vetas, 3950-4160 m, 17.i.1927 (fl), Killip & Smith
21167 (NY).
VENEZUELA. Apuré: Paramo de Pata de Judio, 30 km al S. de San Vicente de
Revancha, 3000-3200 m, 19.i.1968 (fl), Steyermark & Dunsterville 101140 (NY,
VEN). Mérida: Péramo de Los Leones (La Lagunita, La Canada Grande), W. of
MucurubA, 3500 m, 31.v.1930 (fl & fr), Gehringer 138 (F, GH, NY, VEN); Villa de
Caracas, Las Lagunetas, xi. (fl & fr), Moritz 1167 (BM, GH, K, KW, S). Tachira:
Paramo E] Batallén, 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.i.1973 (fr),
Cuatrecasas, Ruiz-Teran & 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 Mérida 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 Belén 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 Mérida, 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 H. pimeleoides can be distinguished from that
of H. cardonae by the broader leaves with flabellate venation visible beneath.
THE GENUS HYPERICUM L.
51. Hypericum caracasanum Willd.
Sp. Pl. 3: 1458 (1802); Choisy, Prodr. monogr. fam. Hypéric. 58 (1821); Kunth in
Humboldt, Bonpland & Kunth, Nova Gen. et Spec. Pl. 5: 186 (1822); Choisy in
DC., Prodr. syst. nat. regni veg. 1: 554 (1824); Turcz. in Bull. Soc. Nat. Moscou 31
(1): 389 (1858); R. Keller in Bull. Herb. Boissier 11, 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.1800 (fl), Humboldt
& Bonpland 649 (B, holotype, F!, US!, photographs; P!, isotype).
Shrub or shrublet 0-5—1-8 m tall, erect, with branches strict, pseudo-dichotomous or
lateral. Stems yellow- to orange-brown, 4—6-lined and compressed when young, the
subfoliar ridges broad, soon or eventually terete, cortex exfoliating in strips;
internodes 1—4 mm long. Leaves subsessile to broadly or narrowly petiolate with
petiole up to 1-5 mm long, + densely imbricate, tetrastichous, ascending to outcurv-
ing, deciduous above petiole without fading; lamina 5—14(-17) x 2-7 mm, +
narrowly elliptic or ovate-elliptic to oblong, plane to incurved or slightly cucullate,
midrib slightly prominent to impressed beneath, margin plane to recurved or slightly
incrassate, paler beneath or concolorous or with margin pinkish, not glaucous,
coriaceous; apex acute to subacute, base angustate, the petioles not or scarcely
sheathing but united to form narrow interfoliar ridge; basal vein 1, with 1—3(4) pairs
of ascending lateral branches, not visibly branching, not impressed beneath, tertiary
reticulation not visible; laminar glands dense, usually visible beneath. Inflorescence
1-flowered, with pseudo-dichotomous branches from node below; pedicel 2-5—5-5
mm long, not or scarcely incrassate upwards; upper leaves not transitional. Flowers
12-20 mm in diam. , stellate. Sepals 5—11 x 2-4-5 mm, broadly or narrowly elliptic or
oblong-elliptic to obovate-spathulate or oblanceolate, acute to subacute; veins 3-7,
sometimes branched distally, with midrib slightly prominent; glands linear, distally
punctiform. Petals bright? to deep yellow, 8-13 x 3-5-6 mm, 1-4-2 x petals,
oblanceolate; apiculus acute; glands mostly linear, distally interrupted to punct-
iform. Stamens c. 50-70, longest c. 4-7 mm long, c. 0-5 X petals. Ovary 2-3-5 x 2
mm, ovoid-ellipsoid; styles 3, 4-5 mm long, c. 1-5—2-:5 x ovary, diverging and
sometimes incurved; stigmas narrow or clavate. Capsule (5)6-8 x (2)3-4 mm,
cylindric-ellipsoid, equalling sepals. Seeds c. 1-2 mm long, ecarinate; testa finely
scalariform.
In humid or shaded habitats in the paramo and subparamo; (1600) 2000-2765 m.
Venezuela (Distrito Federal, Miranda, Aragua, Anzoategui, Sucre). Map 20 (p. 87).
H. caracasanum and H. cardonae are vicarious species which differ in leaf shape,
thickness and glandularity, and in stigma shape, but H. caracasanum is more
restricted in morphological variation and distribution than is its vicariad. In H.
caracasanum the leaf glands are always visible and the stigma is not or scarcely
enlarged. It appears to be directly related to H. jaramilloi.
The extreme easternmost population of H. caracasanum, in the Cerro de Turumi-
quire (Sucre), has smaller, more crowded leaves and somewhat smaller flowers than
the rest of the species, thus showing a trend in morphology as well as geography
towards H. ekmanii from Hispaniola. The variation between the Sucre population
and the rest is apparently almost continuous, and therefore subspecies is the
appropriate rank for the two populations.
51a. H. caracasanum subsp. caracasanum
Brathys caracasana (Willd.) Spach, Hist. nat. vég. Phan. 5: 448 (1836) in Annls Sci.
nat. (Bot.) II, 5: 366 (1836).
Icones: Steyerm. & Huber, Fi. 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
/
v
U
89
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, Anzodtegui).
VENEZUELA. Anzoategui: Rio Mantiales E. of Bergantia, Cerro Peonia above
Santa Cruz, 2350 m, 21.iii.1945 (fl), Steyermark 61667 (F, NY, S, VEN). Aragua:
prope Colonia Tovar, 1854-1855 (fl), Fendler 40 (GH, K, MO, P). Distrito Federal:
Silla de Caracas, Cordillera de Avila, between Los Venados and Pico Oriental,
1675-2640 m, 27—28.11.1944 (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 1mm; styles 4-5 mm long, 2-2-5 x ovary.
On steep north-facing sandstone slopes in subparamo vegetation; 2360-2910 m.
Venezuela (Sucre). Confined to the Cerro Turumiquire.
VENEZUELA. Sucre: Cerro de Turumiquire, 1925 (fl), Tate 329 (US).
52. Hypericum ekmanii A. H. Liogier
in Phytologia 47: 183 (1980). Type: Dominican Republic, Peravia, San José de Ocoa,
2000 m, 4.vi.1978 (fl & fr), A. & P. Liogier 27672 (SDM, holotype; GH!, isotype).
Icon: Fig. 13B.
Shrub up to 0-75 m tall, erect, forming thickets, with branches ascending, pseudo-
dichotomous and sometimes lateral. Stems orange-brown, 4-lined and compressed
when young, the subfoliar ridges broad, soon 2-lined, eventually terete, cortex
exfoliating at first in strips between subfoliar ridges; internodes 2—4 mm long. Leaves
with short petiole (0-5—1 mm long), densely spreading or imbricate, tetrastichous,
deciduous above petiole without fading; lamina 6-10 x 2—4 mm, elliptic, plane to
slightly concave, scarcely cucullate, midrib not or slightly prominent beneath,
margin incurved, not incrassate, concolorous, not glaucous or slightly so beneath,
subcoriaceous; apex acuminate to apiculate, base angustate to cuneate, the petioles
not sheathing but united to form very narrow interfoliar ridge; basal vein 1, with 1
short near-basal pair of branches or unbranched; laminar glands dense, visible on
both sides. /nflorescence 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 xX sepals, oblong-oblanceolate; apiculus
subacute; glands nearly all punctiform. Stamens c. 80, longest 5—6 mm long, c. 0-6 x
petals. Ovary 2 X 1 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 4%; 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 José de Ocoa, S. slope of
Tetero de Majia, c. 2550 m, 1.ii1.1929 (fl & fr), Ekman H. 11718 (S, US).
Ekman (on the back of the label) states that the species is quite common in ‘these
mountains’ (i.e. Sierra de Ocoa) and occurs at 1300 m near Bejucal. The gap in
distribution between H. ekmanii (Hispaniola) and its nearest relative, H. caracasa-
num subsp. turumiquirense (NE. Venezuela), is likely to have resulted from long-
distance dispersal and therefore cannot be adduced to support the theory that North
and South America were originally linked via the Greater Antilles and the Yucatan
Peninsula (see p. 9).
IZ
NORMAN K. B. ROBSON
Dyes.
Map 21 52. H. ekmanii @ ;53. H. pycnophyllum A ;54. H. millefolium @.
53. Hypericum pycnophyllum Urban
Symb. Anitill. 7: 523 (1913). Type: Dominican Republic, La Vega, Loma Rosilla,
1700 m, 6.vii.1912 (fl), Fuertes 1749 (By, holotype; NY!, P!, W!, isotypes).
Icon: Fig. 13C.
Shrub 0-2—1 m tall, erect, with branches very strict, pseudo-dichotomous and lateral.
Stems reddish-brown, 2-lined and compressed when young, the subfoliar ridges
broad, very soon terete, appearing articulated when old, cortex exfoliating at first in
strips between subfoliar ridges; internodes 1-1-5 mm long. Leaves petiolate with
short petiole (c. 0-5 mm long), densely imbricate, markedly tetrastichous, deciduous
above petiole without fading or with attached cortex below after fading; lamina 3—9
x 0-5-2 mm, very narrowly elliptic to linear, incurved to canaliculate, + cucullate,
midrib impressed beneath, margin narrowly hyaline, dull above, lucent beneath, not
glaucous, coriaceous; apex acute, base angustate to above ‘articulation’ then
broadening, pairs forming narrow interfoliar ridge; basal vein 1, unbranched;
laminar glands dense, obscure, visible above only. Inflorescence 1-flowered, terminal
and on lateral branches, sometimes with single or pseudo-dichotomous branches
from node below; pedicel absent or up to 3 mm long, not or slightly incrassate
upwards; upper leaves not transitional. Flowers 10-20 mm in diam., stellate to
obconic. Sepals 5—6(—7) X 1-2 mm, subequal, lanceolate to oblong-lanceolate,
acute to acuminate, cucullate, with margin narrowly hyaline; veins 3—7, unbranched,
midrib not prominent; glands linear, uninterrupted. Petals deep? yellow to orange-
yellow, 7-14 x 3-5-7 mm, c. 1-6 X sepals, oblanceolate; apiculus acute; glands
linear, distally interrupted. Stamens 55, longest 4-5—6-5 mm long, c. 0-6 X petals.
Ovary c. 2 X 1 mm, ovoid; styles 3, 3-5—4 mm long, 1-5-2 X ovary, diverging-
incurved; stigmas subclavate. Capsule 5—7 X 2-5-3 mm, narrowly ovoid, shorter
than sepals. Seeds c. 1 mm long, ecarinate; testa finely ribbed-scalariform.
In open Pinus occidentalis forest and among rocks at higher altitudes; 1700-3175 m.
Dominican Republic (Santiago, La Vega, San Juan, Azua). Map 21.
DOMINICAN REPUBLIC. Azua: El Pico del Yaque, 3125 m, 3.i.1944 (st),
Jiménez 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.1976 (fl), A. & P. Liogier 25040
(NY). Santiago: Monci6n, top of Cerro Lucio, c. 2525 m, 12.vi.1929 (fl & fr), Ekman
H.12850 (C, F, K, NY, S, US).
The specimens from higher altitudes have shorter leaves. Hypericum pycnophyllum
is apparently more widespread than H. ekmanii and links it morphologically with H.
millefolium.
THE GENUS HYPERICUM L.
54. Hypericum millefolium Urban & Ekman
in Ark. Bot. 20 A, no. 15: 79 (1926). Type: Haiti, Massif de Selle, Morne de la Selle,
Morne Emérillon, 2000 m, 1.1925 (fl & fr), Ekman H. 3126 (B7, holotype; K!,
NY!, S!, isotypes).
Icon: Fig. 13D.
Shrub up to c. 2 m tall, erect, juniperoid, with branches strict, pseudo-dichotomous
(extension) and shorter, lateral (flowering). Stems 4—6-lined and ancipitous when
young, persistently 4-angled and ancipitous until cortex exfoliates at first in strips
between subfoliar ridges, then terete, fragile, green becoming pale grey; internodes
1-2 mm long. Leaves sessile, imbricate to curved-ascending, tetrastichous, eventu-
ally deciduous with attached cortex below after (?) fading; lamina 0-7-2 x 0-8-1
mm, lanceolate-triangular, incurved, markedly cucullate, midrib impressed _ be-
neath, margin narrowly hyaline, dull above, lucent or + glaucous beneath,
coriaceous; apex acute, pungent, base broad, subamplexicaul, pairs united to form
narrow 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
linear, uninterrupted. Petals deep? yellow, 6-7 x 2-5-3 mm, 2 X sepals, oblan-
ceolate; apiculus very acute; glands linear, distally subpunctiform. Stamens c. 50,
longest 4—5 mm long, 0-6—0-7 x petals. Ovary 1-5 x 1 mm, broadly ovoid; styles 3,
4—5 mm long, c. 3 X ovary, diverging-incurved; stigmas narrow. Capsule 4 x 2mm,
ovoid, exceeding sepals. Seeds not seen.
Among steep limestone rocks; 1750—2350 m.
Haiti (Massif de la Selle). Map 21.
HAITI. Ouest: 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), Pétionville, top
of Morne Cabaio, 2350 m, 10.iv.1927 (fl), Ekman H. 7979 (C, S).
Hypericum millefolium, with its ericoid habit, is at first glance quite distinct from any
other species in sect. Brathys. Closer investigation, however, shows it to be similar to
the smaller form of H. pycnophyllum. Its triangular leaves and numerous fragile
flowering stems, nevertheless, make it easily recognizable.
55. Hypericum ruscoides Cuatrec.
in Ciencia Mex. 4: 63 (1943). Type: Colombia, Putumayo, Paramo de San Antonio
del Bordoncillo, between El Encano and Sibundoy, 3250 m, 4.1.1941 (fl), Cuatre-
casas 11720 (COL, holotype; F!, P!, US!, isotypes).
Shrub 0-1-1-5 m tall, erect or with stems sometimes decumbent and rooting at the
base, with branches strict, pseudo-dichotomous and sometimes lateral. Stems yel-
low-brown, 4-lined and compressed when young, the subfoliar ridges broad, even-
tually terete, cortex exfoliating in strips between subfoliar ridges; internodes 2-5-6
mm long. Leaves petiolate with petiole 1-5-3-5 mm long, ascending to spreading,
tetrastichous, deciduous above petiole without fading; lamina 8-18 x 3—-6(—8) mm,
narrowly ovate or lanceolate to narrowly oblong-elliptic or the uppermost elliptic,
plane or slightly conduplicate, not or rarely carinate, slightly cucullate, midrib not or
slightly impressed beneath, margin not distinct or rarely subincrassate, concolorous,
sometimes ferrugineous, + lucent or dull and undulate-papillose on both sides,
sometimes glaucous, coriaceous; apex acute to apiculate, base angustate, the
petioles not sheathing but united to form narrow interfoliar ridge; basal vein 1, with
2-4 pairs of obscure ascending branches, rarely visibly branched or reticulating;
laminar glands dense, sometimes visible on either or both sides. Inflorescence
1-flowered, with pseudo-dichotomous branches from node below; pedicel 4-8 mm
03
94
NORMAN K. B. ROBSON
long; upper leaves often broader but not transitional. Flowers 15-25 mm in diam.,
stellate. Sepals 5-10 x 1-5—4 mm, obovate-spathulate to narrowly oblong (unequal
to subequal), acute to obtuse; veins 5, branched or not, midrib not prominent; glands
linear, distally punctiform. Petals deep yellow to orange-yellow, 8-15 X 4-6 mm, c.
1-5 X sepals, obovate-spathulate; apiculus subacute to obtuse; glands linear, distally
punctiform. Stamens 30-60, longest 4—6-5 long, c. 0-5 x petals. Ovary 2-3 x 1-5-2
mm, ovoid; styles 3, 2-5—4 mm long, c. 1:3 X ovary, suberect to divergent; stigmas
broadly capitate. Capsule 4-8 x 3-3-5 mm, cylindric-ellipsoid to ellipsoid, shorter
than sepals. Seeds c. 1-5 mm long, ecarinate; testa finely scalariform.
In paramo; 2450-3800 m.
Colombia (Cordilleras Oriental, Central and Occidental from Boyaca and Cundina-
marca to Putumayo), Ecuador (Pichincha and Napo: Volcan Cayambe). Map 22.
COLOMBIA. Boyaca: Villa de Leiva, Parque Igaque, Quebrada Carrizal, 3400 m,
25.xi.1984 (fl & fr), Wood 4621 (K). Caldas: Navarco, 3000 m, 12.ix.1951 (fi),
Romero-Castaneda 3021 (COL). Cauca: Cordillera Central, Paramo de las Moras,
3800-3700 m, 19.ii.1969 (fl), Cuatrecasas 27424 (BM, COL, US). Cundinamarca:
Municipio de Fomeque, paramo de Chingaza, cerros vecinos a la Laguna y limites
con el Depto. del Meta, 3300 m, 2.xi.1966 (fl), Huertas & Camargo 6572 (COL).
Huila: Cordillera Oriental, 20 km SE. of Gigante, 20.ix.1944, Little 8687 (COL).
Meta: Paéramo de Sumapaz, Cerro Nevado de Sumapaz, 3725 m, 29.i.1972 (fl), Cleef
1206 (BM, COL, U). Narifio: Pasto a Sibundoy, c. 3000 m, 5.i.1952 (fl), Fernandez &
Knoth 1023 (COL). Putumayo: Valle de Sibundoy, Sibundoy, 2800 m, 30.vi.1953
(fl), Schultes & Cabrera 20138 (GH, NY). Tolima: Cordillera Central, Municipio de
Map 22 55. H. ruscoides A; 60, H. selaginella O.
THE GENUS HYPERICUM L.
Roncevalles, paramo de Normandia, 3200 m, 8.xi.1980 (fl), Camargo 7508 (COL).
Valle: Cordillera Occidental, Los Farallones, cerro Alto de Buey, 3500-3600 m,
11.x.1944, Cuatrecasas 17917 (BM, F, US), 17940 (COL).
ECUADOR. Pichincha: Cordillera Oriental, Laguna de Chiqui, NE. of
Cayambe, 3800 m, 4.vi.1944 (fl), Acosta Solis 8151 (F). Napo: ENE. of Cayambe
Mtn, 3210 m, 6.xii.1961, 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.
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 Aztcar, 3800-3900 m,
15.v.1982 (fl), Dllgaard, Holm-Nielsen et al. 38464 (AAU!, holotype).
Shrub 0-4-2 m tall, erect to decumbent but not rooting, with branches strict,
pseudo-dichotomous or lateral. Stems orange-brown, 4-lined and ancipitous when
young, eventually terete, cortex exfoliating in irregular flakes; internodes 1-2 mm
long. Leaves sessile or rarely subsessile, erect to narrowly spreading, + imbricate-
tetrastichous, deciduous above the base after fading; lamina 7-13 x 1-5-4 mm,
narrowly elliptic to linear-oblong, plane or slightly incurved, scarcely to markedly
cucullate, midrib impressed beneath, margin not or very narrowly hyaline, concol-
orous, rather glaucous, + coriaceous; apex acute, base angustate, not sheathing,
pairs united to form rather shallow interfoliar ridge; basal vein 1, (always?) unbran-
ched; laminar glands dense, visible above and rarely also beneath; marginal glands
rather dense. Inflorescence 1-flowered, with pseudo-dichotomous branches from
node below; pedicel 3—4 mm long; upper leaves not transitional. Flowers c. 20-25
mm in diam., stellate. Sepals 7-5-9 X 1-8—2-3 mm, oblong-oblanceolate, acute, not
or slightly incurved-cucullate; veins 5, branched, not or scarcely prominent beneath;
glands striiform to punctiform. Petals (bright?) yellow, 12-16 x 4-10 mm, c. 1-5—1-8
X sepals, oblong-oblanceolate; apiculus acute; glands linear to striiform. Stamens c.
80-90, longest 6-10 mm long, 0-5—0-7 x petals. Ovary c. 3 X 2-5 mm, broadly ovoid;
styles 3, 3-6—4-3 mm long, 1-:2—1-4 Xx 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
95
NORMAN K. B. ROBSON
Map 23 56. H. llanganaticum O; 57. H. struthiolifolium @.
essentially only in the narrower leaves with a slightly cucullate apex and narrower
stigmas. The remaining specimens are nearer to H. struthiolifolium but differ from it
in having more crowded tetrastichous leaves (which are dull blue-green or glaucous,
not lucent) and unribbed, oblanceolate to oblong sepals.
57. Hypericum struthiolifolium Juss.
in Annls Mus. Hist. nat. Paris 3: 161, t. 16, f.2 (1804) [‘struthiolaefolium’|; Choisy,
Prodr. monogr. fam. Hypéric.: 58 (1821), in DC., Prodr. syst. nat. regni veg. 1: 553
(1824); R. Keller in Bull. Herb. Boissier II, 8: 176 (1908), in Engler & Prantl, Nat.
Pflanzenfam. 2nd ed. 21: 181 (1925). Type: Peru, without precise locality or date
(fl & fr), Dombey s.n. (P-JUS!, holotype; F!, P!, isotypes).
Brathys_ struthiolifolia (Juss.) Spach, Hist. nat. vég. Phan. 5: 449 (1836)
[‘struthiolaefolia’|, in Annls Sci. nat. I, 5: 366 (1836).
H. struthiolifolium var. genuinum Triana & Planchon in Annis 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, I, 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 2093
(B+, holotype; F!, GH!, NY!, US!, photographs; G!, isotype).
Shrub up to c. 0-5 m tall, erect or sometimes decumbent and rooting, with branches
strict, pseudo-dichotomous or sometimes lateral. Stems orange-brown, 4-lined and
ancipitous when young, eventually terete, cortex exfoliating in irregular flakes;
internodes 2—7 mm long. Leaves sessile, suberect to spreading or slightly outcurving,
not or scarcely imbricate, deciduous above the base after fading; lamina 8-15 x
1-5-2-5 mm, very narrowly elliptic to linear-oblong, + incurved- cucullate, midrib
THE GENUS HYPERICUM L.
mostly impressed beneath, margin not or slightly hyaline, concolorous, lucent, not
glaucous, + coriaceous; apex acicular to subacute, base angulate, not sheathing,
pairs united to form shallow interfoliar ridge; basal vein 1, occasionally with 1-2
pairs of obscure branches; laminar glands rather dense, visible above but not
beneath. /nflorescence 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 5S—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 Piscohuafuna, 2700 m, viii.1938 (fl),
Sandeman 239 (K); Bagua, Cordillera Colan E. of La Peca, 2880 m, 8.viii.1978 (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: Chilifruta to Huancayo, 11
km, 3700 m, 30.x.1968 (fl), B. & C. Maguire 61657 (BM, NY). Pasco: Oxapampa,
Cordillera Yanachaga 12 km SE. of Oxapampa, 2700-2800, 7.x.1982 (fl), Foster
9048 (BM, MO).
H. struthiolifolium was one of the first species in sect. Brathys to be described, and
the name has frequently been misapplied. The species is confined to north and
central Peru, where it is apparently relatively uncommon. It differs from H.
llanganaticum in the less crowded, more incurved, lucent (not glaucous) leaves and
the usually smaller flowers with sepals that are ribbed and usually broader below the
middle. Also, its decumbent stems sometimes root. Its distribution overlaps con-
siderably that of the derivative H. andinum, but these species remain distinct.
58. Hypericum andinum Gleason
in Torreya 29: 137 (1929). Type: Bolivia, Mt. Tunare, 1891 (fl & fr), Bang 1026 (NY!,
holotype; BM!, F!, GH!, K!, MO!, US!, W!, isotypes).
H. struthiolifolium var. parviflorum R. Keller in Bull. Herb. Boissier I1, 8: 182 (1908)
[‘struthiolaefolium’|. Type: Bolivia, La Paz, Larecaja, Majabaga, Catarguata,
2800-3200 m, x.1858—v.1859 (fl & fr), Mandon 790 (G!, holotype; BM!, GH!, K!,
NY!, P!, S!, W!, isotypes).
Shrub or shrublet 0-1-0-5 m tall, erect or decumbent and rooting, bushy with
branches strict, pseudo-dichotomous or lateral. Stems reddish to yellowish-brown,
4-angled when young, soon 2-lined then terete, cortex exfoliating in irregular flakes;
internodes 1-4 mm long. Leaves sessile, suberect to outcurved, subimbricate to
spreading, deciduous above the base after fading; lamina 3-5—-10 x 0-8-1-:5 mm,
narrowly oblong-elliptic to linear, incurved-cucullate, midrib mostly impressed
beneath, margin + narrowly hyaline, otherwise concolorous, not glaucous, sub-
coriaceous to chartaceous; apex acute to rounded, base cuneate to parallel, not
sheathing, pairs united to form shallow interfoliar ridge; basal vein 1, unbranched;
laminar glands dense, impressed, visible beneath. Inflorescence 1-flowered, with
pseudo-dichotomous branches from node below; pedicel 3—6 mm long; upper leaves
not transitional. Flowers 10-18 mm in diam., stellate. Sepals 3-5-7 x 0-8-2 mm,
broadly to narrowly lanceolate, subacuminate to acute, cucullate, with hyaline
margin; veins (3)5(7), unbranched, + prominent; glands linear, punctiform, in distal
third. Petals golden yellow, 6-10 x 3-5 mm, c. 1-5 X sepals, narrow obovate to
obovate-oblong; apiculus acute to obtuse or obsolete; glands shortly striiform to
punctiform. Stamens 20-50, longest 3-6 mm long, c. 0-5 x petals. Ovary 1-8—2-3
Ae
o7
98
NORMAN K. B. ROBSON
(—3:3) X 1-1-2 mm, narrowly ovoid; styles 3, 1-5—2-5 mm long, 0-6—0-7 xX ovary,
free, spreading; stigmas broadly capitate. Capsule 4-6 x 1-2-3 mm, ovoid, shorter
than sepals. Seeds not seen.
On open or stony grassy slopes and Sphagnum bogs, sometimes near the snowline;
2400-4200 m.
Peru (La Libertad, Junin, Huancavelica, Cuzco, Puno), Bolivia (La Paz, Cochabam-
ba, Santa Cruz). Map 24.
PERU. Cuzco: Paucartambo Valley, 26 km from Cuzco to Pilcopata, 3425 m,
27.ix.1968 (fl), B. & C. Maguire 61564 (BM, NY). Huancavelica: Tayacaja, between
Colcabamba and Paucarbamba, Arriba de la Hda. Jocas, 3900 m, 20.iv.1954, Tovar
2020 (NY). Junin: Huancayo, 11.1939 (fl), Soukup 389 (F). La Libertad: Huamachu-
co, Jalca de Huaguil, 3950 m, 24.vi.1958 (fl), Lopez & Sagastegui 2736 (NY). Pum:
Ayapata, vi.1854 (fl & fr), Lechler 1817 (K, P, W).
BOLIVIA. Cochabamba: Road to Chapare [from Cochabamba], Km 104, 3100
m, 23.xii.1926, Steinbach 650 (F, MO, NY,S, U, US, WIS). La Paz: Sorata, 3900 m,
ii.1886 (fl & fr), Rusby 1350 (MICH, NY). Santa Cruz: Comarapa, Cerro de San
Mateo, 3400 m, 25.x.1928 (fl & fr), Steinbach 8535 (BM, F, GH, K, MO, NY, S, VU).
Although the area of H. andinum overlaps considerably that of H. struthiolifolium,
the former can be distinguished by the shorter capitate styles and the narrower,
usually less acute, recurving leaves with markedly hyaline margins.
59. Hypericum lancioides Cuatrec.
in Ciencia Mex. 4: 65 (1943). Type: Colombia, Putumayo, Paramo de San Antonio
del Bordoncillo, between El Encano and Sibundoy, 3250 m, 4.i.1941, Cuatrecasas
11721 (COL, holotype; F!, US!, isotypes).
Shrub c. 0-1-1 m tall, erect and bushy or + decumbent and caespitose, branches
strict, pseudo-dichotomous and lateral, or spreading (very rarely pinnate). Stems
orange-brown, 6-lined and compressed when young, the subfoliar ridges broad,
eventually terete, cortex splitting between and below subfoliar ridges; internodes
0-5—4(—6*) mm. Leaves sessile or rarely narrowing to a pseudopetiole up to 2 mm
long, ascending to subimbricate, sometimes tetrastichous, deciduous above base (or
pseudopetiole) after twisting spirally; lamina 7-15 x 1—-1-4(—4*) mm, very narrowly
elliptic-oblong to linear incurved to canaliculate or rarely subconduplicate, +
markedly cucullate, midrib impressed beneath, margin distinct, narrowly to rather
broader hyaline, concolorous, lucent beneath, undulate-subpapillose above,
glaucous or not, coriaceous to somewhat succulent; apex acute, base angustate to
parallel-sided, not or scarcely sheathing, pairs united to form narrow interfoliar
* 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 1(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 (S—)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—S mm long, c. 0-5 X
petals. Ovary 1-5-2-5 x 0-8-1-5 mm, ellipsoid to subglobose; styles 3(4), 1—2(—3*)
mm long, 0-5—0-9(—-1-1*) x ovary, outcurved to suberect; stigmas broadly capitate.
Capsule 4-5 x 1-5-2:5 mm, ovoid-ellipsoid to cylindric-ellipsoid, shorter than
sepals. Seeds 0-8-1 mm long, ecarinate; testa finely scalariform.
In damp (sometimes marshy) or sheltered areas of the open paramo; 2870-4700 m.
Western Venezuela (Mérida), Colombia (Santandér to Narifo) 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
Mérida and has lateral flowers crowded below the terminal flower. Intermediate
specimens between the subspecies occur in Cauca.
59a. Hypericum lancioides Cuatrec. subsp. lancioides
H. struthiolifolium var. genuinum Triana & Planchon in Anals Sci. nat. (Bot.) IV, 18:
293 (1862) [‘struthiolaefolium’] pro parte, quoad spec. cit.
Map 25 59. H. lancioides a. subsp. lancioides A, b. subsp. congestiflorum Ne
ao
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
Yo; b-f X 3; i-m X 4). A. B. & C. Maguire 61808; B. Cuatrecasas et al. 28120; C. Barclay & Juajibioy
7407.
H. strictum sensu Gleason in Bull. Torrey 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-
THE GENUS HYPERICUM L.
developed branches from lower nodes. Sepals oblong to linear, with veins promin-
ent.
3000-4100 m.
Southern Colombia (Cundinamarca and Meta southwards) and Ecuador (Carchi to
Cotopaxi).
COLOMBIA. Cauca: Cordillera Central, Paramo de las Moras, 3800-3700 m,
12.xii.1969 (fl & fr), Cuatrecasas 27419 (BM, COL, US). Cundinamarca: Cordillera
Oriental S. of Usmé, paramo de Chisaca above Laguna Grande and L. Negra,
9-11.xi.1958 (fl), Barclay & Juajibioy 6191 (COL, MO, NY). Huila: Municipio de
La Plata, Inspeccién de Policia de Santa Leticia, Paramo de La Candelaria, 2370 m,
15.vii.1975 (fl), Diaz, Lozano & Torres 595 (COL). Narifio: 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.1946, Schultes & Villareal 7960 (COL, F).
Putumayo: Pasto to Sibundoy, Paramo de San Antonio, 2790-2880 m, 13.iii.1953 (fl
& fr), Schultes & Cabrera 18828 (COL, GH, NY, U). Valle: Cordillera Occidental,
Los Farrallones, cerro de La Torre, La Laguna, 3500-3550 m, 1.viii.1946 (fi),
Cuatrecasas 21870 (BM, F, P, U, US).
ECUADOR. Carchi: Tulcan to Maldonado, c. 43 km, SW. slopes of Volcan
Chilas, 3740 m, 6.viii.1976 (fl), Ollgaard & Balslev 8549 (AAU, BM). Cotopaxi:
Ruminahui, 3850 m, 17.ii.1947 (fl & fr), Aubert de La Rue s.n. (P). Imbabura: Lago
San Marcos, Cayambe, 3360 m, 26.xi.1961 (fl), Cazalet & Pennington 5349 (K).
Napos: Laguna Porjuantag, ENE. of Volcan Cayambe, 3750 m, 10.1i.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.1944 (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] in Annls 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, Mérida,
Sierra Nevada de Mérida, 2870 m, 1842 (fl), Linden 432 (COL?, holotype; P!,
isotype).
Icon: Fig. 14B.
Shrub 0-1-0-5 m tall, erect and bushy or decumbent and caespitose with branches
strict, lateral (very rarely pinnate), spreading. Leaves sessile, ascending to imbricate,
tetrastichous, linear. Inflorescence branches elongate, sympodial (from terminal
node) and very short, lateral, with flowers solitary or in triads in a congested
spiciform thyrse, immediately below terminal flower or with several intervening
sterile nodes. Sepals oblong-lanceolate, with veins not or slightly prominent.
2870-4500 m.
Western Venezuela (Mérida), north-eastern and central Colombia (Santandér to
Cauca).
COLOMBIA. Arauca: Sierra Nevada del Cocuy, Quebrada El Playon, Plan de S.
José, 3625 m, 9.vi.1973 (fl), Cleef 10051 (U). Boyaca: Paramo de Chita, cabeceras
del Rio Casanare, Hoya Laguna Chorros Blancos, 3340 m, 24.v.1973 (fl), Cleef
9931A (COL, U). Caldas: Paramo del Quindio, 4100-4300 m, 15-20. viii.1922 (fl),
Pennell & Hazen 9874 (GH, K, S). Cauca: Paramo de Purace, 3500 m, 26.v.1944 (fl),
Killip & Lehmann 38548 (COL). Cundinamarca: Paramo de Palacio, c. 600 m al
NW. de Mina de Cal, 3520 m, 13.v.1972 (fl), Cleef 3733 (BM, COL, U); Lagunas de
Buitrago, 3555 m, 27.iv.1973, Cleef 9588A (COL). Meta: Paramo de Sumapaz,
Cerro Nevado de Sumapaz, 3615 m, 29.i.1972, Cleef 1269 (COL, U). Risaralda:
Municipio de Pereira, alredores de la Laguna Otién, 4000-4300 m, 3.ii.1980 (fl),
Diaz, Valencia & Jaromillo 1754 (COL). Santandér: Paramo del Almorzadero,
101
102
NORMAN K. B. ROBSON
3500-3700 m, 20 vii.1940 (fl), Cuatrecasas & Garcia Barriga 9998 (COL, F). Tolima:
Paramo de Herves, Eresno, 3600 m, 14.vii.1939 (fl), Hanbury-Tracy 603 (K).
VENEZUELA. Mérida: Paéramo de Timotes, 3200 m, 25.i.1939 (fl), Alston 6616
(BM, NY); Quebrada de Saisay, 3880 m, 10.iv.1930, 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,
11-29.iv.1959 (fl & fr), Barclay & Juajibioy 7407 (COL!, holotype; MO!, NY!,
isotypes).
H. selaginoides N. Robson in Cleef, Veg. Param. Colomb. Cord. Or.: 305 (1981),
nomen.
Icon: Fig. 14C.
Shrublet up to 0-15 m tall, prostrate or ascending, forming tufts up to 150 mm in
diam., with branches pinnate, spreading. Stems orange-brown, 6-lined and com-
pressed when young, the subfoliar ridges broad, eventually terete, cortex splitting
between and below subfoliar ridges; internodes c. 0-5 mm long. Leaves sessile,
densely to loosely imbricate, tetrastichous, deciduous above base or more usually
only with the cortex after fading; lamina 4-7 x 0-5—0-8 mm, linear, incurved-
canaliculate, cucullate, midrib impressed beneath, margin broadly hyaline, con-
colorous, sublucent beneath, sublucent to dull above, sometimes subglaucous,
coriaceous; apex acute to subacute, base parallel-sided, not sheathing, pairs united
to form narrow interfoliar ridge; basal vein 1, unbranched; laminar glands dense
above, rather sparse or absent beneath. Inflorescence 1-flowered, with pseudo-
dichotomous branches from node below and also short congested lateral branches
(racemiform); pedicel absent or up to 2 mm long; upper leaves not transitional.
Flowers c. 8 mm in diam., stellate. Sepals 3-4 x 0-8-1-2 mm, oblong to triangular-
lanceolate (equal), acute, cucullate, margin broadly hyaline; veins 3(5?), unbran-
ched, slightly prominent; glands mostly linear. Petals bright yellow, tinged red
outside, 4-5-6 X 2-2-5 mm, 1:5 X sepals, obovate-oblong; apiculus very short or
almost absent; glands linear, distally punctiform. Stamens c. 13, longest 3-4 mm
long, c. 0-65 X petals. Ovary 1-5—2-5 x 1-5-2 mm, subglobose; styles (3)4—5, 0-8-1
mm long, 0-4—0-5 x ovary, outcurved; stigmas broadly capitate. Capsule c. 3 x 3
mm, cylindric-globose to globose, shorter than sepals. Seeds 0-7—0-8 mm long,
ecarinate; testa finely scalariform.
In dry and stony or dampish paramo or superparamo; 3300—4340 m.
Colombia (Boyacé, Arauca, Cundinamarca), in the Sierra Nevada del Cocuy,
Pdramo 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.1973 (fl & fr), Cleef
8944 A (COL). Boyaca: Sierra Nevada del Cocuy, Valle de los Corallitos, c. 4300 m,
31.vii.1957 (fl), Grubb, Curry & Fernandez-Perez 178 (COL, K, US); Paramo de
Pisva, Socha to La Punta Km 61.5, 6 km al NE. de Los Pinos, 3600 m, 10.vi.1972 (fl &
fr), Cleef 4328 (BM, COL, U). Cundinamarca: Macizo de Sumapaz, Cuchilla La
Rabona, 3900-3950 m, 17.vii.1981 (fl), Diaz et al. 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
Mérida, which are somewhat intermediate in habit and size of parts.
THE GENUS HYPERICUM L. 103
61. Hypericum cymobrathys N. Robson, sp. nov.
H. terrae-firmae Sprague & Riley affinis, sed habitu humiliori, foliis brevioribus
crassioribus, interdum leviter cucullatis, supra basin deciduis, floribus minoribus in
cymes 3-12-floris dispositis, 3-stylis, inter alia differt. Type: Colombia, Boyaca, La
Uvita, by road from Chita, 3000-3050 m, 16.ix.1969 (fl), Cuatrecasas & Rodriguez
27805 (COL!, holotype; BM!, US!, isotypes).
Shrub 0-5—1 m tall, erect, with branches + strict, pseudo-dichotomous or occasional-
ly lateral. Stems orange-brown to purplish-brown, 4-lined and markedly ancipitous
when young, soon 4-angled, cortex exfoliating in irregular flakes; internodes 2-5 mm
long. Leaves sessile, spreading from above base, tetrastichous, deciduous from
above base without fading; lamina 9-18 x 3-8 mm, oblong to elliptic, plane, slightly
cucullate, midrib prominent towards the base beneath, concolorous, glaucous,
coriaceous; apex acute to obtuse, base cuneate to angustate, sheathing, saccate,
pairs united to form narrow interfoliar ridge; basal veins 5—7, subparallel, apparently
unbranched, tertiary reticulum not visible; laminar glands dense, prominent to +
impressed. Inflorescence 3—12-flowered, crowded (monochasial)?, with pseudo-
dichotomous branches from node below; peduncle and pedicels 3-5 mm long,
sometimes incrassate upwards; upper leaves intermediate, bracts narrowly elliptic-
oblong, acute. Flowers c. 15 mm in diam., stellate or cyathiform. Sepals 6-8 x
2:2-2:4mm, 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 2 x 1-5 mm, broadly
ellipsoid; styles 3, 3 mm long, 1-5 < 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); Péramo 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, 1.1976
(fr), Aguirre & Rangel 391 (COL).
Although the cymose inflorescence of H. cymobrathys is reminiscent of H. mexica-
num, its flowers and leaves are very different. Its relationships appear to be with H.
terrae-firmae, although the much smaller flowers, 3—12-flowered inflorescence, and
smaller leaves make it superficially unlike it. Indeed, it appears to be an isolated
relict species at the base of a clade that includes H. chamaemyrtus, H. gnidioides, and
the H. gentianoides group, as well as H. denticulatum and its relatives (see part 8).
3. Acknowledgements
I am especially grateful to Dr Bassett Maguire (NY), who has helped my studies of tropical American
Hypericum in many ways; I have also received valuable assistance from Dr Antoine Cleef (U), Dr José
Cuatrecasas (US), Dr Richard Howard (A), Dr Alicia Lourteig (P), Dr James Luteyn (NY), Dr Benjamin
@llgaard (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 actuacién 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. Jn 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.
Hist. (Bot.) 8: 55-226.
1981. Studies in the genus Hypericum L. (Guttiferae). 2. Characters of the genus. Bull. Br. Mus. nat.
— 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 Jiménez, 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. O. 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
(= Sla) 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 var. scherzeri (= 7)
30
caracasanum var. turumiquirense
Steyerm. (= 51b) 90
cardonae Cuatrec. (50) 87*
carinosum R. Keller (9) 33, 37*
cassiopiforme N. Robson (29)
et aaa
chamaemyrtus sensu Gleason
(= 4) 26
costaricense N. Robson (38) 70,
wi
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
(1), 3° 325
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
lancioides sensu Steyerm. (= 45)
80
lancioides subsp. congestiflorum
(Triana & Planchon) N.
Robson (59b) 99*, 100*, 101
lancioides subsp. lancioides (59a)
99* , 100*
lancioides forma pygmaea
Cuatrec. (59a) 100
laricifolium Juss. (20) 47, 48*,
49*
laricifolium sensu Bentham
(=35) 67
laricifolium var. acerosum
(Kunth) Weddell (= 20) 47
laricifolium var. glaucum R.
Keller (= 48b) 85
laricoides Gleason (= 20) 47
lindenii R. Keller (= 19) 45
lindenii sensu Cuatrec. (= 15) 40
llanganaticum N. Robson (56)
95, 96*
loxense Bentham (18) 43
loxense subsp. aequatoriale (R.
Keller) N. Robson (18a) 44*
loxense var. aequatoriale R.
Keller (= 18a) 44
loxense subsp. loxense (18b) 44*,
45
THE GENUS HYPERICUM L.
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)
TT.
marahuacanum subsp.
chimantaicum N. Robson (44c)
71*, 78*, 80
marahuacanum subsp.
marahuacanum (44a) 71*, 78*,
79
marahuacanum subsp.
strictissimum N. Robson (44b)
TAA 1 8o 509
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)
Paget Pt
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,
1
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
pimelevides Planchon & Linden
ex Triana & Planchon (31) 60,
O2c
piriai (6) 28, 29*
platypetalum Turcz. (= 20) 47
platyphyllum Gleason (= 3c) 25
platyphyllum Gleason pro parte
(= 25) 55
prietoi N. Robson (28) 57*, 58
prostratum Cuatrec. (42) 75, 77*
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 Diels (= 17) 41
ruscoides Cuatrec. (55) 93,
94*
105
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. B 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
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
NORMAN K. B. ROBSON
terrae-firmae Sprague & Riley
(1) 18, 19*, 21*
tetrastichum Cuatrec. (47) 82*
thuyoides Kunth (13) 37*
thymifolium Cuatrec. (= 3a) 23
thymifolium sensu Planchon &
Triana pro parte (= 9) 33
thymifolium sensu Turcz. (= 19)
45
trianae N. Robson (= 19) 45
valleanum N. Robson (33) 62*,
64, 66*
weberbaueri R. Keller (= 57) 96
weberbaueri sensu Gleason
(= 45) 80
wurdackii N. Robson (37) 62*,
69
Studies in the genus Hypericum L. (Guttiferae)
N. K. B. Robson
When completed in nine parts, this series of papers will constitute a monograph of
Hypericum, St John’s wort. This ancient genus, with an almost world-wide distribu-
tion, is of great scientific interest, as well as of considerable importance in horticul-
ture and, to a lesser extent, pharmacy. Parts 1 (1977) and 2 (1981) are introductory,
and the detailed taxonomy begins with part 3 (1985), in which the first seven sections
(sects 1—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. 0 565 08003 2. £28.80
Titles to be published in Volume 16
Studies in the genus Hypericum L. (Guttiferae)
7. Section 29. Brathys (part 1)
By N. K. B. Robson
The lichen genus Ramalina in Australia
By G.N. Stevens
An annotated list of vascular plants collected in the valleys south of
Mt Everest
By G. Miehe
An illustrated catalogue of type specimens in the Greville diatom herbarium
By D. M. Williams
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd, Dorchester
Bulletin of the
British Museum (Natural History)
The lichen genus Ramalina in Australia
G. Nell Stevens
Botany series Vol 16 No 2 25 June 1987
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British Museum (Natural His rae us OF
Cromwell Road
London SW7 5BD Issued 25 June 1987
The lichen genus Ramalina in Australia
G. Nell Stevens
Department of Botany, University of Queensland, St Lucia, 4067, Australia
Contents
DOOD fee esssiacsoses ah tee talent ia nena Peet rds a earn aecck ne RN A
MIVT OAC SNOD erg sc s4e. caine coco uence das noe vuneimnce rend cat ee eR ew aaa cece tice
Historical understanding of Ramalina and Ramalinaceae ....................0..02.
Collectors of Australian Ramalinat ss... cccects recs Gekcdetnnosdeedimawasaever tees soe
PAALOTIGIS AIRNINOTNOOS: occscrscthersscse mata oO ee enc rele deen
PROS aaa ies ae coe ved eo wapads Pee coats ena PE nee es eR eis een cho
PIOLDDOIOD suchas raise cebu daeetel aoac a einem 8. a encom een
PRBUOY 2 Porndevnnar caren ees aL ena dN ea eae eats RG Ah a Reet bone eceay sa aaceahions
COIS stun ocsveteoe riers ecenderes oh ceVas) sau entoalen a eethecar tear
Correlation between secondary-product chemistry and ecogeography
i (he RAMANA SUDITARINEG COMDICK: f.ioi56c4s dadeices ue Aocigs thes caseunneccr abeeeen
Distribution patterns in relation to climatic indices .................0.ccccecceeeeeeees
RORGORLODNY) 0555 cis o5 2 Uo ade nu sacs uecy sda cts ueuiso des oocvace ta cre fal daaashueetasaetienns
POAC OUI MCOREADINY tors csac eve. fscc Ncw ctea adi tass ccave ty anian Witwer igcinetaes tions
EN UN ae AO ROY Ee RP Racer halo eR ET aa Sa Ader a Vee oral re Sa
PROUOTIOMMG RMTIMS taser es cre nies sree ered tees ce ae tie i ri deuincre se wile aekee |
AY ERGI PATON epoch yt ences teeeanneckae cere tech caeas tenet niva titan eita fin aia
Pescripiien of the semis anand. 25 We = cs een yaw ee ee
Enumeration Obthe Australian taxa. pyuccutastusiontucincaersreereneean deities
PCy emmenriire/hd W PNVIS ETAL 8 Se ee ccahn oacties Meueaeets saencrencsemals meen’
di PRE SEDC TINES «ic ace Psd 2 aE asec canteen MOM ae rs tauesuersasoelakekiNes
re ed fy ORR ear eT eee TEN. eT RE PrP aAR aac re Aree me TN
Bios Ro CRPIONICTISUS osc faz pr elatace Pe clade tosh eae Bs sta don MW Oe SRG aaidbre pheeoo tet D bee TTL:
Wee AG MRIRMEN Ci 5 oe vss schions 35 S0ie8 RoNty ent Tet dare Se neck Ea nia dac nna eesy Vane ake
GEM RELCLIUN ES rai sticiere draarsaet che te CaCO aan atg bos cake vise Pete nan tuimnadei oes acces
yee ee POP ROLE re Et ae Bea seOk Coen ar nea a A AT TARE en we es he Pasiese CMM EEL cE
6 C4 ETE Tel) ea RR I fo Pes EE Ns Pr SN a ORME a rR eee
8
9
10
2. Rr
PCR UOVEO a See ae Seer te aye oe ued en ee cate ee ee Pee ae TT ee EAE ice Tes env GEE.
Sede PRCTULVIANG atc tiger cede more aaa ae Pete ae Ak See Feces ere ay Gee LES
POCA Oo PR PO ese 8 eA ode o a Feng Ate heen ea ae Miu)
ef | SER ORE OPO ESR LE AEES SCR TS on ARON So LEONE ier APONTE REE Ronee ee E
—
od .
Dy Po PO 7
ICSE CA ONE GRASS VE RRORER AREY SnaPS SG se PEM Eee Cor egy REROPE Te Petry eae nN OSE De
OPE PCUICR, crete ede er ee eaten eh Moe uae ect aedehae ee eee rac caed is cedonde
MO PoP SUM VOXEL eer ire eos IO o ae eae oi ae wae ate ea cate ET oe ec genau
Ure OPEC tas pases ee sas seeks se nen tia Bs heen eusvidea de ie Aoctsn hee wee vntes aoneeiee
Appendix. Disposition of exluded Ramalina taxa ...............cccccceceenecececueneees
PROB OW ISU CE IOREG rs dass ah cts feces esses ettut ddaes actin ceaynctseaith us vk dyes eaawanienns
Re NC tote es ene cera c iene tae gn seae Se Eat os oe raul sages ka eek aR IONS
MIO ay ee rd ge ea dah py eae AE Co nis Pe eean dk acon dos eee eeI GENIN ad
Bull. Br. Mus. nat. Hist. (Bot.) 16 (2): 107-223
PAWIATEI IGE Sos ote aies cao tata thesia nie ia pen vaso tecsie da Rader oN ALT eS ERE Ae eh
Soy FAL ¢ SURE COCR PARE ROMEY ESTO NERA ROPERSANC ELE (oy RE PRACS Marne ARENT IEA
Issued 25 June 1987
108 G. NELL STEVENS
Synopsis
This paper presents a revision of the lichen genus Ramalina in Australia. Of the 60 Ramalina taxa
previously listed for Australia in catalogues only nine of those names are retained, six with unchanged
status. Twenty-eight taxa belonging to 20 species are recognized for Australia. Two are new species: R.
filicaulis and R. tropica. A new variety R. subfraxinea var. norstictica and a new subspecies R. inflata subsp.
australis are described, and four names, R. celastrisubsp. 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 microtherm) as used to divide the Australian phanerogams. On the basis of species
diversity it is hypothesised that the genus had its origin in west Gondwanaland (South America and Africa)
and then spread to the east. It is suggested that subsequent speciation occurred within Australia.
Introduction
The genus Ramalina has been studied in the northern hemisphere for over two hundred years
and several hundred taxa have been named. In the southern hemisphere very little research has
been undertaken and no revisional work had been carried out on the Australian Ramalinae
before the present study. During the course of previous research into the taxonomy and ecology
of twenty-four lichen genera found on eastern Australian mangroves (Stevens, 1978) it became
apparent how necessary it was for a revision of the Australian representatives of the genus to be
undertaken.
This present work provides a taxonomic revision of the Australian Ramalinae as well as an
ecological survey and a biogeographic assessment of the genus. Research into the Australian
taxa in this genus also contributes to the overall understanding of the biogeography of allied
Ramalina species which occur in the other Gondwanaland regions of Africa, India, and South
America.
The genus is regarded as a difficult one and its morphological plasticity is renowned. Howe
(1913-14) aptly summed up the dilemma when he stated ‘there is little difficulty in recognizing
the characteristics of the genus Ramalina. The species on the other hand, have presented a most
difficult problem, caused largely no doubt by the innumerable intergrades that present them-
selves, and by the unwarranted description of new species’.
This problem was encountered by the author when confronted by the record of 60 different
Ramalina taxa reported for Australia (Wetmore, 1963; Weber & Wetmore, 1972) in catalogues
which listed all taxa recorded for Australia in papers published from 1804 to 1972. It was
apparent quite early in the study that many of these names were incorrect; some were synonyms,
others misapplication of the names of northern hemisphere species which they superficially
resembled.
Other recent mention of Australian Ramalinae occurs in an unpublished key to the genus
Ramalina by Dahl (1971) and in Filson & Rogers (1979), but in both of these accounts little
preliminary research had been carried out before naming the taxa. Most of the existing literature
summarizing Australian material was therefore of little help in the present study.
There are few modern publications recording research which include the southern hemi-
sphere Ramalinae. Krog & Swinscow (1974, 1975, 1976) have published detailed work on the
East African Ramalinae. Rundel (1978) has dealt with the Ramalina usnea complex in North
and South America and in a paper on the lichens of Tristan da Cunha, Jorgensen (1977)
discussed six species of Ramalina. No other publications discuss this genus in detail; however, a
series of papers by Osorio published from 1967 to 1983 list the lichens found in Uruguay,
Paraguay, Brazil, and Argentina and these include some species of Ramalina. An account of the
lichens of New Zealand (Galloway, 1985) which includes the genus Ramalina lists only nine
species.
RAMALINA IN AUSTRALIA 109
During the present research four months were spent in Europe and U.S.A. visiting herbaria
(BM, FH, H, HAW, O, S, UPS and US) and becoming familiar with northern hemisphere
Ramalina species. This enabled an insight to be gained into the reason why Australian species
were often given the names of the northern hemisphere species. The type material of species
which occur in other southern continents was also examined where possible.
In this study the approach has been to reinforce the taxonomic revision of the genus by
incorporating ecological data in order to explain phenotypic variation, and to enable the
delimitation of the taxa to be made more confidently. The extreme morphological variation
within the species was a major problem. The extent of such variation in Australian material had
not previously been gauged, nor was there any conception of the evolutionary relationships that
might exist between the different taxa.
Many thousands of specimens therefore needed to be examined before any decisions could be
made on the delimitation of species. Examination of too few specimens makes it impossible to
understand the breadth of morphological plasticity exhibited by a single species, and the early
taxonomy of Australian Ramalinae was based on single or at most several thalli which were
usually collected by someone other than the taxonomist.
Collection of Ramalina material by the author began in 1975, when the lichens on mangroves
were studied (Stevens, 1978). Since then collecting trips have covered parts of New South
Wales, Victoria, Tasmania, Western Australia, and Northe 1 Territory and innumerable
collecting forays were made in many areas of Queensland. After several expeditions inland it
was found that the genus Ramalina was restricted to a zone varying within 100—200 km of the
coastline, thus eliminating the need to collect in the arid interior (Fig. 1).
Observation of the different ecological habitats occupied by any one species was made to
enable an appreciation of the variation that occurs in the morphology of a species under different
environmental pressures. As well as sampling different habitats in the one region, thalli were
BO
v4,
Py Sa
N Ae
fg}
‘ : on
. ey e Se e °° y
ee as © ay
ate Sats
8
Fig. 1 Ramalina distribution in Australia.
110 G. NELL STEVENS
collected (wherever possible) from latitude to latitude in order to assess if a continuum existed
between the different morphologies found over a wide geographical range.
To a large extent this research has been carried out on personal collections of fresh material,
supplemented by collections made by present day Australian lichenologists, material available
from herbaria in each of the Australian states, and to a far lesser extent on old collections held in
overseas herbaria. The latter proved to be of little use as only vague information concerning the
location of the collection was ever recorded, e.g. New Holland. In contrast the Australian
material was well documented on the whole but the specimens remained unnamed or bore
doubtful identifications.
All of the Australian types were examined and, where possible, compared with fresh material
from the type location. In this way the amount of variation that occurred within the particular
species could be assessed. It was found in some instances that the type represented one extreme
in a large population of morphotypes.
An attempt has been made to show possible evolutionary relationships between some of the
taxa, based on morphological and/or chemical evidence associated with geographical patterns of
distribution, as suggested by Imshaug & Brodo (1966).
This revision has treated all Australian material as belonging to the genus Ramalina Ach., as
observations during the present study did not substantiate separation into the two genera
Ramalina Ach. and Fistulariella Bowler & Rundel. No species belonging to any other divisions
within the Ramalinaceae (Bowler, 1981) occur in Australia. A representative collection has
been deposited in BM.
The Australian environment
Although it is beyond the scope of this study to provide a detailed discussion of the geography,
climate, botany, and geology of Australia, a general outline is presented to enable a better
appreciation of the factors which are influencing Ramalina distribution (Fig. 1).
(1) Geography
The area of Australia is approximately 8,700,000 km?, larger than all of western Europe, and its
immense coastline measures some 36,700 km. In total it comprises the Australian mainland, the
large island of Tasmania, and numerous small islands around its coastline. It is washed by three
oceans, the Pacific, Southern, and Indian, and four seas, the Tasman, Timor, Coral, and
Arafura. It extends from approximately 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 Australian land mass is a vast flat peneplain with an
elevation between 200 m and 500 m. In the north-west area the highest peak in the Hamersley
Ranges exceeds 1200 m in height, and in central Australia the Macdonnell and Musgrave Ranges
reach a similar height. Much of the interior of Australia is arid sand-ridge desert; the arid zone
RAMALINA IN AUSTRALIA Lh
extends to the western and southern coasts between the moist south-west corner of Western
Australia and the more fertile south-east of Australia. Extensive salt lakes occur in the south and
south-western regions of the arid interior.
(2) Climate
The general climatic characteristics of Australia are dictated by its latitudinal position lying
astride the mid-latitude high pressure belt. The prevailing wind patterns and general aridity over
continental Australia reflect the dominating control of eastward-moving anticyclonic cells which
track between 37°—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
Australia and Tasmania (Nix, 1981).
Summer rainfall predominates in the north, winter rainfall in the south, and along the eastern
coast a more uniform distribution of rainfall is common (Figs 2A & B). The eastern coastline of
Australia receives a substantial rainfall, which ranges from over 2000 mm in north Queensland
to 741 mm in Westernport Bay, Victoria. The western coastline receives far less precipitation
per annum, partly due to the lack of any mountain ranges close to the coast. The annual
precipitation ranges between 230 mm and 840 mm per annum with the highest recordings in the
southwest corner where winter rains occur. Winter rainfall is predominant as far north as 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 Australia. 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 Australian land mass receives over 3000 hours of sunshine per year
while the east coast is reported to receive between 2500 and 3000 hours, indicating that
cloudiness is higher near the coast.
The Queensland coast as far south as Mackay and the Western Australian coast as far south as
Carnarvon are subject to tropical cyclones, normally between December and March, which
cause very heavy rainfall and accompanying strong winds. Sea surface temperatures around
Australia range from 15°—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 listed on the top
left hand side of Table 1, representing those which occur in relatively high rainfall areas and have
112 G. NELL STEVENS
[]
GF
| (A)
(B)
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).
e
Life form of
Foliage projective cover of tallest stratum
tallest stratum 100-—70% 70-50% 50-30% 30-10% <10%
Trees >30 m Tall closed- Tall — a —
forest open-forest
Trees 10-30 m Closed- Open-forest Open-forest Woodland Open-
forest woodland
Trees <10m 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 <0:25m — —_— — Dwarfopen- Dwarf open-
heathland heathland
Hummock grasses — —_— — Hummock Open hummock
grassland 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 30m in height, but in monsoonal
northern Australia and coastal Queensland, low closed-forests occur which are composed of
trees reaching less than 10 m in height. During the wet season these forests have a FPC of 100%,
but during the dry winters the FPC may be as low as 70% as some of the trees are semi-
deciduous. In inland Queensland the low closed-forest is the only formation which supports any
Ramalina species.
(b) Tall open-forest (with sclerophyllous understorey with or without tree ferns) occurs in
sub-humid sites of eastern, south-eastern and south-western Australia. Overstorey FPC in such
a community varies from 70% to 50%. The dominant tree species in the overstorey is
Eucalyptus, whereas the low tree/tall shrub stratum (less than 10 m high) contains 18 genera of
angiosperms (Specht, 1981a). Several species of Ramalina have been found growing on twigs in
the understorey, but rarely do species of Ramalina grow on eucalypts.
(c) Open-forest (with either heathy sclerophyll or grassy understorey). This formation is the
most widespread in Australia, occurring in the north, east, and southern parts of the continent;
each area experiences quite different rainfall patterns ranging from summer rains to summer-
winter uniform rainfall, to winter rains respectively. The genus Eucalyptus dominates these
114 G. NELL STEVENS
communities, although species of Casuarina and Acacia are also plentiful. The overstorey FPC
for open-forest communities varies between 70-30%. No Ramalina species grow in the tropical
open-forests, but occur on Casuarina and Acacia in the temperate communities.
(d) Mallee open-scrub (with savannah-chenopod understorey). This structural formation is
common in the drier Mediterranean climate of southern Australia where species of Eucalypt
dominate, but other genera, including Acacia are scattered between the mallee eucalypts.
Overstorey FPC in these areas can be as low as 30%. The relatively low rainfall (250-800 mm
p.a.) occurs in winter, followed by hot dry summers. Four species of Ramalina have been
collected in this formation.
Seral 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 Australia. The structure of such communities varies from
closed-forest to woodland to tall shrubland. The major concentration of mangrove vegetation
(approx. 28 species) is confined to the tropics gradually dwindling in species numbers in the
subtropics and eventually being reduced to one species, Avicennia marina, towards the
temperate zone. Both FPC and the amount of rainfall vary considerably depending on the
latitude of the mangrove community.
Those mangrove species which act as phorophytes to the Ramalinae are Rhizophora stylosa,
Rhizophora apiculata, Ceriops tagal, Bruguiera gymnorhiza, Avicennia marina, Aegiceras
corniculatum, Lumnitzera racemosa, and Excoecaria agallocha.
(4) Geology
The saxicolous species of Ramalina are totally dependent on the presence of rock outcrops
within their distribution range. Corticolous species, however, are only indirectly influenced
through the presence or absence of particular plant communities which grow on the different soil
types.
The four main rock types involved in the areas where Ramalina species occur are (1) basaltic
rocks (2) metamorphics and fine grained sedimentary rocks (3) sandstones and (4) granites.
The major areas of basalt are Tertiary in age, and are confined to the Eastern Highlands. Most
of these elevated areas are forested with either rain-forest or tall open-forest. Additional areas
of basalt of Quaternary age are found in north-east and central-east Queensland, supporting low
closed-forests, and western Victoria at lower altitudes on the coastal plain which supports
extensive areas of open-woodland and grassland.
In eastern Australia the metamorphic rocks and many fine-grained, compacted sedimentary
rocks which are associated with them are of Palaeozoic age, and have been folded and elevated,
forming mountain ranges and hilly country in the Eastern Highlands. They have been intruded
by granite batholiths, which also make up large areas, having been revealed by prolonged
erosion e.g. New England, Murrumbidgee, Kosciusko, and many other masses in the eastern
states. These are areas of high elevation subject to cold winters.
The sandstones which are most important are those sub-horizontal deposits of Mesozoic age
RAMALINA IN AUSTRALIA 1i5
forming for example, much of the surface of the Sydney Basin, Moreton Basin, and the
Carnarvon Range. The Grampians are composed of dipping sandstones of Palaeozoic age.
In South Australia and Western Australia only a few species of Ramalina occur on trees
growing on soils derived from old granites and metamorphic rocks (schists and quartzites).
There are very few occurrences of Ramalinae in areas of calcareous rocks of the Tertiary basins
and Pleistocene dune deposits.
Historical understanding of Ramalina and Ramalinaceae
(1) THE GENUS RAMALINA
The genus Ramalina was first described by Acharius (1810) when he distinguished it from
Parmelia by its cartilaginous thallus and apothecial characters.
Acharius (1810) listed ten species (together with their forms) as belonging to the genus,
namely Ramalina homalea, R. linearis, R. complanata, R. peruviana, R. polymorpha, R.
fraxinea, R. fastigiata, R. scopulorum, R. farinacea, and R. pollinaria. Except for the first, all of
these species have been retained in the genus to the present day.
Many more taxa have been added to this original number in the last 175 years, some being
removed later into separate genera because of differences found in the anatomy, the spore shape
and size, or the colour of pycnidia. Montagne (1852) examined the cortex of R. homalea Ach.
and found that the hyphae were arranged perpendicular to the surface without any mechanical
support tissue in the cortex; he therefore erected a new genus Desmazieria Mont. to incorporate
all Ramalina species possessing this anatomical feature, using R. homalea as the type.
Massalongo (1854) erected the genus Cenozosia Massal. based on R. inanis Mont. as the type
and characterized by its fistulose spongy thallus with a single layer of branching cells in the
cortex. This monotypic genus has been retained as a distinct taxon up to the present.
By using the colour of spermatia, as well as anatomical and thallus differences, Stizenberger
(1862) divided the genus Ramalina into three parts (i) Cenozosia (Massal.) (type R. inanis Mont.)
as a species with thallus hollow, spermatia black (ii) Desmazieria (Mont.) (type R. ceruchis Ach.)
for those species with simple cortex, thallus interior cottony and spermatia black and (iii)
Euramalina Stizenb. (type R. scopulorum Retz.) for species having a double cortex, thallus
interior cottony, and spermatia pale or colourless.
Nylander (1870) also used colour of spermogonia conceptacles for initial division within the
genus, separating species with (1) totally black spermatia (2) partially black or (3) pale or
colourless spermatia. Within group (1) he then used differences in anatomical structure of the
cortex to separate R. inanis Mont. (= Cenozosia Massal.) from the rest. He placed five other
species in another group which corresponds to Desmazieria Mont., namely R. ceruchis Ach., R.
combeiodes Nyl., R. homalea Ach., R. flaccescens Nyl., and R. testudinaria Nyl.
Ramalina melanothrix Laurer was separated from the rest because of its two-layered cortex
(= Euramalina Stizenb.); its present position is in the genus Trichoramalina Rundel & Bowler.
Group (2) contained only R. carpathica K6rber, 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-
sae Hue.
Steiner (1904) based his divisions of Ramalina on the arrangement of cortical tissue external to
116 G. NELL STEVENS
the algal layer. His section Corticatae (outer layer consisting of more or less transversely
arranged conglutinated hyphae) was equivalent to Desmazieria Mont., his section Bitectae
(outer layer as above overlying a tissue of longitudinally arranged conglutinated hyphae) was
similar to Euramalina Stizenb. He made a third division Ecorticatae (cortex composed of a layer
of longitudinally oriented conglutinated hyphae which formed a closed ring). Ramalina arabum
(Ach.) Meyen & Flotow is the type for Ecorticatae. Choisy (1954) commented that this thallus
structure was similar to Alectoria Ach.
Howe (1913-14) used spore characters to distinguish the sections he created within the genus
Ramalina as he thought this represented a more natural classification. Section Ellipsosporae R.
H. Howe contained species with ellipsoid or oblong spores (9-20 x 3:5—7-5 zm) and section
Fusisporae R. H. Howe contained species with long fusiform spores (16-35 xX 3-7 wm) 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 Tenuicorticatae R. H. Howe and section Bitectae Steiner.
Section Tenuicorticatae contained species without a chondroid element and this section he
further divided into subsection Solidae Du Rietz (species with a solid thallus, containing R.
evernioides, etc.) and subsection Tubulosae Du Rietz (species with inflated fistulose thallus: R.
inanis Mont. = Cenozosia Massal.).
Du Rietz maintained section Bitectae Steiner (= Euramalina Stizenb.) but divided it into two
subsections (a) subsection Myelopoea Vainio thallus solid, and (b) subsection Fistularia Vainio
thallus inflated, fistulose. These extra subsections make the classification complicated although
the basic criteria used for division are sound.
Zahlbruckner (1907) combined the sections of both Stizenberger (1862) and Steiner (1904)
and made three divisions in the genus Ramalina: (i) section Ecorticatae Steiner (ii) section
Corticatae Steiner (which incorporated Cenozosia Massal. and Desmazieria Stizenb.) and (iii)
section Euramalina Stizenb. (= Bitectae Steiner). Zahlbruckner subdivided Euramalina
Stizenb. using the Vainio names Fistularia Vainio and Myelopoea Vainio and series Teretius-
culae Vainio and Compressiusculae Vainio.
Present day taxonomists are still divided in their acceptance of which species should be placed
in the genus Ramalina sens. str. Bowler & Rundel (1977) established the new genus Fistulariella
Bowler & Rundel for those species originally placed in Ramalina stirps Fistularia Vainio. They
placed emphasis on the hollow, usually perforate thallus as traits warranting genus status, yet
Australian material belonging to stirps Fistularia Vainio present a broad range of morphology
varying from almost totally compressed to totally fistulose. It is therefore regarded here that the
trait of inflation of the thallus should have no greater emphasis placed upon it than the roundness
of branches.
Rundel & Bowler (1978) proposed a new generic name Niebla to replace the genus name
Desmazieria Mont. as they found the former name was a homonym for the earlier described
genus Desmazeria Dumortier (Poaceae), and 13 taxa were transferred by them from Desma-
zieria and Ramalina into Niebla Rundel & Bowler.
Of the various divisions put forward up to the present day, the taxonomic system used by
Vainio would most suitably classify the Australian material, e.g. the Australian Ramalinae
would broadly divide into three groups: (1) fistulose thalli (2) terete and solid thalli and (3)
compressed and solid thalli.
(2) THE FAMILY RAMALINACEAE
Agardh (1821: 93) erected the family Ramalinaceae [‘Ramalinez’], in which he placed the genus
Ramalina. Since that time the genus has been alternatively included with Usnea and other
RAMALINA IN AUSTRALIA Ly
fruticose genera in the family Usneaceae, or isolated in the family Ramalinaceae. Table 2 shows
the various classifications which record Ramalina in Ramalinaceae (with or without other
members of Usneaceae) and the diversity of opinions as to the position of the genus.
Apparently Zahlbruckner (1907) and Smith (1921) did not regard septate spores nor a cortex
with mechanical tissue sufficiently different to separate Ramalina from other genera in the
family Usneaceae; but Watson (1929) regarded spore septation as most important and placed
Ramalina in the Ramalinaceae.
With modern day improvement in techniques of microscopy and chemistry together with a
better understanding of the fungal/algal composition, the natural relationships which exist
between the genera are being more fully understood.
Follmann & Huneck (1969) placed three genera in the family Ramalinaceae based on their
chemotaxonomy and anatomy (viz. Ramalina, Ramalinopsis, and Desmazieria). Poelt (1974)
also regarded these three genera as forming a natural entity and placed them in Ramalinaceae,
but he added a fourth genus which was not closely related, e.g. Speerschneidera. Culberson &
Culberson (1970) pointed out that chemical evidence supported segregation of Ramalinaceae as
a separate family as no species in the Usneaceae proper produces orcinol meta-depsides, a
category of substances richly represented in Ramalina.
Henssen & Jahns (1973) have kept the genus Ramalina in the family Ramalinaceae, because
of its two-celled spores and exobasidial conidiophores, and because of the specific lichen
substances it contains. They stated that the structure of the apothecium of Ramalina resembles
that of the Parmeliaceae in that there is a more or less complete algal layer beneath the
hymenium, but the Ramalinaceae differ from the characteristic cupular exciple of the
Parmeliaceae in that the algal layer is next to, not separated from the loosely interwoven
subhymenium.
Keuk (1979) divided the genera in Ramalinaceae into two groups based on anatomical and
chemical differences, and colour of pycnidial walls (a) Ramalina, Ramalinopsis, and Tricho-
ramalina and (b) Niebla (= Desmazieria Mont. and Cenozosia Massal.).
The most modern division within the family Ramalinaceae has been based on cortical
diversity. Bowler (1981) defined seven genera in the Ramalinaceae: Ramalina Ach., Fistu-
lariella Bowler & Rundel, Niebla Rundel & Bowler, Ramalinopsis (Zahlbr.) Follm. & Huneck,
Trichoramalina Rundel & Bowler, Dievernia M. Choisy, and Cenozosia Massal. Of these, the
genus Ramalina contains the largest number of taxa.
It can be seen from Table 2 that all recent authors who have specifically dealt with the question
of relationships of genera within the two families Ramalinaceae and Usneaceae, have con-
sidered them to be distinct morphologically, chemically, and taxonomically.
Collectors of Australian Ramalinae
Because the early explorers in Australian waters sailed in the temperate latitudes, it was only to
be expected that the first Ramalina collected in Australia would be a temperate region species;
this collection was made by Labillardiére, a botanist on the French ship ‘Recherche’ which sailed
along the Great Australian Bight and visited Tasmania in 1791 (Ducker, 1979). This first
specimen, held at H, (H-NYL 37226) was annotated ‘Ramalina leiodea Nyl. Labillardiere, C.
van Dieman’. This was an erroneous identification which has now been determined as R. inflata
J. D. Hook. & Taylor.
In 1802, Robert Brown in his voyages with Matthew Flinders collected R. inflata from Port
Jackson (Sydney, N.S.W.), but this specimen was determined as Lichen fastigiatus Ach. by
Crombie (1880).
When J. D. Hooker accompanied Sir James Ross in his Antarctic expedition in the ships
‘Erebus’ and ‘Terror’ in 1839, he collected the type of R. inflata J. D. Hook. & Taylor on the
Lord Auckland Islands and from New Zealand he collected the type of R. 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 Ny]l. as a new species but
118
G. NELL STEVENS
Table2 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 Ramalina, Alectoria, Collema
Order Secundus Peltati 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 Ramalina, Alectoria, Evernia,
Dufourea, Dactylina
Tr. Usneei Usnea, Neuropogon, Chlorea
Stizenberger, E. 1862 Class Tr. Usneaceae
Homothalami Ramalineae Ramalina, Dactylina, Cetraria,
Evernia, Dufourea
Usneaceae Usnea, Alectoria, Oropogon,
Argopsis
Vainio, E. A. 1890 Order Tr. Parmelieae Ramalina, Usnea, Alectoria,
Cyclocarpeae Evernia
Hue, A. M. 1901 Series Radiatae
Cyclocarpae Tr. Ramlineae Ramalina, Dufourea
Tr. Usneae Usnea
Tr. Alectoreae Alectoria
Zahlbruckner, A. 1907. Order Usneaceae Ramalina, Usnea, Alectoria,
Cyclocarpeae Oropogon, Dufourea, Evernia,
Everniopsis, Endocena,
Letharia, Thamnolia,
Siphula, Dactylina.
Smith, A. L. 1921 subseries Usneaceae Ramalina, Usnea, Alectoria,
Cyclocarpineae Evernia, Letharia,
Everniopsis, Oropogon,
Thamnolia, Siphula, Dactylina,
Dufourea, Endocena
Watson, W. 1929 Order Ramalinaceae Ramalina
Parmeliales Usneaceae Usnea, Alectoria, Letharia,
Evernia, Everniopsis, Dactylina,
Dufourea, Thamnolia, Siphula,
Endocena
Rasanen, V.J.P.B. 1943 Order Usneaceae Ramalina, Usnea, Alectoria,
Cyclocarpeae 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. & 1969 Ramalinaceae Ramalina, Ramalinopsis,
Huneck, S. Desmazieria
Henssen, A. & 1973 Order Ramalinaceae Ramalina
Jahns, H. M. Lecanorales
suborder
Lecanorineae
Poelt, J. 1974 Order Ramalinaceae Ramalina, Ramalinopsis,
Lecanorales Desmaziera, Speerschneidera,
suborder Usneaceae Usnea, Alectoria, Bryopogon,
Lecaniorieae Cornicularia, Evernia,
Everniopsis, Himantormia,
Letharia, Sulcaria, Neuropogon,
Oropogon
Bowler, P. A. 1981 Ramalinaceae Ramalina, Fistulariella,
Trichoramalina, Niebla,
Dievernia, Cenozosia,
Ramalinopsis
examination of this material during the present study has shown this specimen to be yet another
morphotype of the very variable R. inflata.
The Reverend Dr. W. Woolls, a school master at Parramatta, N.S.W., in 1832, collected in
that area, and one Ramalina specimen called R. gracilis Nyl. (MEL 9432) is actually R. exiguella
Stirton.
During the period 1842—46, J. P. Verreaux (a French ornithologist) who was sent to Australia
by the Museum of Natural History in Paris, collected widely in Tasmania and mainland
Australia. In his botanical collections were several Ramalina species which he had obtained
during that period, either by collecting them himself or by purchase of material from others
(Lamy, in litt.). Because of this practice of obtaining specimens from other collectors there is
uncertainty about the locality given by Nylander for two ‘type’ specimens he records as collected
by Verreaux from Swan River, Western Australia, these are R. australiensis Nyl. and R.
confirmata Nyl. Both of these species grow in eastern Australia, and the Western Australian
types are the only material collected from that area. Nylander himself appeared unsure of the
locality of R. australiensis as he noted: ‘in Australia (prope Swan River, ni fallor) legit Verreaux
1846’. Verreaux collections held at PC indicate he travelled north as far as Moreton Bay,
Queensland, and collected several Ramalina species from mangroves.
The first woman recorded as a lichen collector was Amalie Dietrich, who collected lichens in
Queensland for a wealthy Hamburg merchant named Godeffroy, for his private museum. Some
of these collections are now housed at Minchen. 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 Mill. 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 R. inflata var. fissa Mill. Arg. from King Island, Bass Strait.
The type of R. lacerata Mill. 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. myrioclada Mill. Arg. from Twofold Bay, New South Wales.
Charles Knight, a New Zealand surgeon, collected lichens in the Sydney area; he named one
specimen R. subgeniculata C. Knight, which was subsequently changed to R. knightiana Zahlbr.
Knight also named R. minuscula var. alba a specimen collected by James Keys at Mt Perry,
Queensland. This name was published by Bailey (1886), but no description was given and later
John Shirley (1888) described it.
Messrs. Pentzke and Hartmann were two collectors in Queensland for the Victorian Botanical
Department (Wilson, 1889). Pentzke collected the type material of R. farinacea var. nervulosa
Mill. Arg. from Daintree River, north Queensland, and C. H. Hartman of Toowoomba
collected the type material of R. geniculata var. compacta Mill. 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 Miller [Arg.] for identification (Wilson, 1889). It was Stirton who named the two
Ramalina types R. exiguella Stirton and R. perpusilla Stirton. Miller Arg. described a
Rockhampton specimen collected by Mrs Thozet as R. farinacea var. squarrosa Mill. Arg. The
type of R. farinacea var. dendroides Mill. 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 R. calicaris var. australica Rasanen
from Barwon Heads. Wilson himself described several taxa of Ramalina, e.g. R. unilateralis F.
Wilson, R. brevis F. Wilson, and R. brevis var. brevissima F. Wilson.
More recent collectors of Ramalinae in Australia are J. H. Willis, Assistant Government
Botanist at the National Herbarium Melbourne (Filson, 1976), A. C. Beauglehole, who
collected Ramalinae in South Australia and Victoria, J. S. Whinray, who made an extensive
survey of the lichen flora of the Bass Strait Islands, and G. C. Bratt and J. A. Cashin, whose
collections of Tasmanian lichens have proved invaluable in the present study.
Other present day lichenologists who have collected Ramalina specimens which have been
used in the present study are A. Archer (N.S.W.), J. A. Elix (A.C.T.), R. Filson (Vic.), G.
Kantvilas (Tas.), R. W. Rogers (Qld.), N. Sammy (W.A.), C. Scarlett (Qld), and R. Seppelt
(S.A.).
Materials and methods
The source of material for this research was threefold. (a) Personal collections (numbering over
10,000 specimens), the majority from Australia, with small collections from the Pacific Islands of
Fiji, New Caledonia, Rarotonga, Tahiti, New Hebrides, New Zealand, and Hawaii. (b)
Herbarium material from institutes, as acknowledged. (c) Herbarium and living material from
private collections, as acknowledged.
Most of the specimens examined during this study were tested by thin-layer chromatography
using the techniques of Culberson (1972). Solvents B and C proved to be the most useful solvents
to separate the acids found in the various species of Ramalina. The chemistry of type material
was also checked. Examination of the surface features of the thallus (e.g. pseudocyphellae,
soralia, and soredia shapes) and spore size and shape, was carried out by use of the Scanning
Electron Microscope (SEM) using either a Cambridge or a Phillips 505. In order to observe the
shape and thickness of the hyphae composing the outer cortex and the supportive tissue thin
sections were examined by TEM, using an AEI Corinth 275 operated at 60 kV. Preparation of
thallus material involved fixation of the tissue in glutaraldehyde, post fixation in osmium
tetroxide, dehydration and then infiltration and embedding with Spurrs medium. The results
were not always successful as the tissue tended to pull away from the medium along the outer
cortex junction.
RAMALINA IN AUSTRALIA 1A!
All climatic data were derived from Meterological Bureau records (Climatic averages of
Australia, 1975).
Results
Morphology
(a) Soralia and soredia: Soralia are defined herein as those areas of thallus where clusters of
soredia are found, and the term soredia is applied to the more or less spherical bodies composed
of algae surrounded by short hyphae which act as vegetative diaspores. These vegetative
reproduction bodies are produced by 11 of the Australian Ramalinae, and their constancy in
occurrence enables this attribute to be used taxonomically.
The position and form of the soralia are also useful criteria in the separation of each sorediate
taxon. Round or ellipsoid soralia are produced by R. pacifica, R. caespitella, R. nervulosa var.
nervulosa, R. nervulosa var. luciae, and R. nervulosa var. dumeticola (Plate 1, fig. 3); these occur
marginally and laminally. Ramalina peruviana and R. tenella produce small, punctiform soralia
(Plate 1, figs 1, 2) which are mainly apical in R. tenella and marginal or lateral in R. peruviana.
The soralia found in the other four taxa are produced in several ways, e.g. (i) by disintegration of
the lower cortex (R. fimbriata, Plate 1, fig. 5) (ii) by the partial separation, both marginally and
apically, of the upper and lower cortices (R. canariensis) (iii) by lack of a continuous lower
cortex (R. reducta, Plate 1, fig. 4) and (iv) by the splitting or disintegration of parts of the lower
cortex at intervals along the branches (R. unilateralis, Plate 1, fig. 6). The soralia terminology
used by Du Rietz (1924) and Beltman (1978) was not generally applicable to the Ramalinae.
The composition of the soredia was examined under SEM; the algal cells were not discernable
but the fragments of hyphae which make up the outer surface of the soredium were observed to
differ in form in several of the taxa. The northern hemisphere species R. capitata is regarded as
having soredia covered by an ‘epicortex-like’ layer (James, pers. comm.) (Plate 2, fig. 1), and
such a structure was looked for in the Australian sorediate taxa. Four species were found to
possess some semblance of an outer covering (Plate 2, figs 2,3, 4,5). Whether this layer should
be compared to the ‘epicortex’ which occurs in some taxa in the Parmeliaceae as described by
Hale (1973, 1981) is debatable; but it can be likened to the ‘Kittsubstanz’ (cementing substance)
mentioned by Peveling (1970). Such a layer may act as a protection against wetting of the
soredia.
The Australian taxa which produce soredia with this outer covering are R. caespitella, R.
reducta, R. tenella, and R. fimbriata, and even in these only a minority of the soredia show it
clearly. The soredia of R. caespitella are small, spherical, and have a partially smooth surface
(Plate 2, fig. 2). Soredia of R. reducta are irregular in shape and size but fusion of the external
hyphal segments was apparent (Plate 2, fig. 3). Ramalina tenella soredia are small but well
defined, and some have a distinct outer covering (Plate 2, fig. 4). Ramalina fimbriata soredia are
Plate 1 (overleaf) SEM photographs showing soralia shapes in some of the Australian Ramalinae. Fig. 1
Apical, punctiform soralia — R. tenella. Scale 10 mm = 530 wm (x 19). Fig. 2 Marginal/lateral,
punctiform soralia— R. peruviana. Scale 10 mm = 150 wm (x 65). Fig. 3 Round and ellipsoid soralia— R.
nervulosa var. dumeticola. Scale 10 mm = 220 wm (xX 44). Fig. 4 Exposed lower surface with chondroid
strands of cortical tissue across area of soralia— R. reducta. Scale 10 mm = 270 «um (X 36). Fig. 5 Patches
of exposed medulla by disintegration of the under side — R. fimbriata. Scale 10 mm = 200 wm (xX 51). Fig.
6 Lower surface split apart and eroded at intervals — R. unilateralis. Scale 10 mm = 530 wm (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 wm (xX 440). Fig. 2 R. caespitella — showing portion of soredia covered by a smooth
layer. Scale 10 mm = 22 wm (x 440). Fig. 3 R. reducta — showing soredia with smooth surface (x 364).
Fig. 4 R. tenella — showing thin layer partially enclosing the soredium (xX 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).
G. NELL STEVENS
Plate 1
RAMALINA IN AUSTRALIA
Plate 2
124 G. NELL STEVENS
large and spherical, and the formation of an external layer could be discerned on many of the
soredia at various stages of development; the most advanced layer formation is shown in Plate 2,
fig. 5. All of these taxa are regarded as having granular soredia.
The soredia produced by R. unilateralis (Plate 2, fig. 6) and by R. canariensis (Plate 2, fig. 8)
are small in size and are composed of both thick and thin pieces of hyphae; there is no external
fusion of the hyphae so the outer surface remains loosely interwoven giving a woolly appearance
to the soredia (Plate 2, fig. 6). The soredia are farinose. The four taxa which belong in the R.
farinacea complex (R. pacifica, R. nervulosa var. nervulosa, R. nervulosa var. luciae, and R.
nervulosa var. dumeticola; Plate 2, fig. 7; Plate 3, figs 1, 2, 3) also produce soredia which have an
irregular surface made up of separate hyphal pieces which are not fused (Plate 3, fig. 2), and the
soredia are farinose. Ramalina peruviana produces soredia which have an irregular surface
(Plate 3, fig. 4).
Fibrils often arise from mounds of farinose soredia (Plate 1, fig. 2) and these features have the
same structure as the soredia (Plate 3, fig. 7).
(b) Apothecia: The production of apothecia as a sexual means of reproduction occurs in all but
four of the Australian Ramalinae (i.e. seven of the 11 sorediate taxa also produce apothecia, but
only rarely, and often the mature ascospores are few and difficult to find). The esorediate taxa
generally produce numerous apothecia.
The position of the apothecia on the branches can be of taxonomic importance, being
terminal, subterminal, marginal, lateral, or laminal. Terminal refers to the apothecia at the end
of a branch. Subterminal refers to apothecia positioned near the end of the branch, with the
branch extending beyond the apothecium, forming a short attenuate branchlet or spur, which is
usually bent at an angle in relation to the main branch. If the spur grows long it makes the branch
appear geniculate. Marginal refers to apothecia on the edges of compressed branches, and
laminal refers to apothecia occurring on the surface of the thallus. If branches are subterete to
terete the apothecia occur laterally. When laminal apothecia occur specifically at the base of
bifurcating branches they are referred to as positioned at the axil of the branch.
Apothecial shape varies from concave to plane to convex; in some species the apothecia
remain concave to maturity, in others the apothecia are concave to plane when immature, but
are markedly convex at maturity. The thalline margin is usually distinct and entire, occasionally
becoming incised at maturity; when the disc becomes convex the margin is often obscured.
The size of the apothecia varies greatly from 0-2 mm to 5-0 mm with exceptions to 10 mm.
Most apothecia are shortly stalked, but sessile apothecia are found in R. australiensis and R.
filicaulis. The colour of the disc is usually yellow-green and pruinose; however, taxa in the R.
celastri complex have been found with orange coloured discs.
The apothecial tissue comprises a colourless hymenium formed of asci and paraphyses. The
asci are clavate to subcylindrical and lie amongst sparsely branched paraphyses over a developed
hypothecium. This hypothecial layer is usually brownish-yellow as is the epithecium [as used in a
lichenological context — Henssen & Jahns (1973)].
(c) Spores: The number of spores per ascus is eight, usually distichous in arrangement (Plate 5,
fig. 5), hyaline, 1-septate, varying considerably in size and shape, being broadly ellipsoidal to
fusiform and either straight to slightly curved or reniform (Plates 4 & 5).
Within a single ascus spores can vary from straight to curved, a feature also noted by Krog &
Plate 3 SEM photographs showing soredia forms and surface features in some of the Australian
Ramalinae. Scale indicated by black bar. Fig. 1 R. nervulosa var. luciae — showing soredia with irregular
surface (x 750). Fig. 2 R. nervulosa var. nervulosa — showing fragments of hyphae making up the outer
surface, without a surface layer (x 945). Fig. 3 R. pacifica — showing soredia with an irregular surface (x
525). Fig. 4 R. peruviana — showing soredium with an irregular surface (x 1300). Fig. 5 Punctiform
pseudocyphellae — R. subfraxinea var. 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 wm (xX 341). Fig. 8
Tuberculate pseudocyphellae — R. subfraxinea var. subfraxinea. Scale 10 mm = 30 wm (X 341).
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Swinscow (1975) and Landr6n (1972) (Plate 4, fig. 4); therefore the shape of spores was not used
as a taxonomic character. The size of spores has also been found to be of limited taxonomic value
as most mature spores of the Ramalina species lie within the size range (8—)10-16 x 3-5-6 um
(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 4um 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 jm 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, 1983a: figs 1B-D, 2A-B). In the
Ramalina subfraxinea complex in Australia some of the taxa produce tubercles resembling
raised warts, the tips of these structures containing small pseudocyphellae (Plate 3, fig. 8). The
term ‘tuberculate pseudocyphellae’ was used by Krog & Swinscow (1975) to describe this form
of pseudocyphellae. Similar surface features occur in several North American taxa, but are
referred to as papillae, as in R. sinaloensis (Bowler & Rundel, 1972b) and R. complanata
(Landr6n, 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, 1983a).
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).
Plate4 SEM photographs showing spore shapes in the Australian Ramalinae. Fig. 1 R. subfraxinea var.
leiodea— single spore. Scale 10 mm = 1-2 wm (X 8000). Fig. 2 R. australiensis spores. Scale 10 mm = 44m
(x 2400). Fig. 3 R. pacifica spores. Scale 10 mm = 4 wm (X 2400). Fig. 4 R. subfraxinea var. confirmata
spores. Scale 10 mm = 4 wm (X 2400). Fig. 5 R. subfraxinea var. norstictica spores. Scale 10 mm = 4 wm
(Xx 2400). Fig. 6 R. subfraxinea var. subfraxinea spores. Scale 10 mm = 4 wm (Xx 2400).
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RAMALINA IN AUSTRALIA 129
1) Cortex
ae Australian taxa the outer cortical layer is usually 20 wm in thickness and is composed of
thick walled, transverse or randomly oriented hyphae; it may be indistinct or even absent. The
inner layer serves as supportive tissue and varies in thickness. It is composed of bundles of more
or less periclinally arranged thick walled hyphae which make up a continuous or interrupted
sheath around the internal medullary hyphae.
The detail in the outer cortex is often difficult to detect under light microscopy when yellowish
granules are present; a problem pointed out by Bowler (1981). To obtain as much detail as
possible, sections were examined under TEM. Both cortical layers of hyphae are highly
gelatinized and a uniform distribution of lumina is apparent; each lumen is surrounded by a thick
matrix of fibrillar material arranged concentrically (Plate 6, fig. 1). Peveling (1974) commented
that it is difficult to determine whether the different layers around hyphae are part of the proper
cell wall or if they are extracellular additions to the wall.
The change in orientation of the hyphae is apparent under TEM. In transverse section the
randomly orientated outer layer hyphae appear elongate whereas the longitudinally oriented
hyphae appear circular (Plate 6, fig. 2). Scattered between the hyphae are outlines of crystals,
which are more dense in the outer region and may be usnic acid crystals (Plate 6, fig. 3).
(a) Anatomical structure as a means of separating taxa. Krog & Osthagen (1980) divided the
species of Ramalina found in the Canary Islands into four anatomical types based on the position
of the supportive tissue: the farinacea type, the decipiens type, the bourgaeana type, and the
duriaei type. The latter two types occur in species now placed in the genera Niebla and Dievernia
and are therefore not relevant to Australian material.
The majority of the Australian Ramalinae could be classed as belonging to the farinacea type
(Figs 3A, B), as they possess a thin outer layer of tissue and a continuous inner layer of
mechanical tissue of uneven thickness. Variations can occur in the arrangement of this inner
supportive tissue if it becomes partly discontinuous, leaving sections of outer tissue unsupported
(Fig. 3C). However, this is not comparable with the ‘decipiens type’ where some of the strands
anastomose across the medulla. None of the Australian taxa could be separated on this trait.
Bowler (1981) pointed out that the Ramalina taxa could be divided arbitrarily into two groups
based on the degree of specialization in the cortical tissue. One group with a narrow, indistinct
outer cortex intergrading with the underlying supportive tissue and the other group with a
distinctly double-structured cortex, but he commented that intergradation could be seen
between the two forms.
The Australian taxa can be divided in this way generally, but thalli within one taxon can often
show differences in thickness of the outer cortical layer. Those taxa with a sparse or indistinct
outer cortex are R. caespitella, R. celastricomplex, R. pacifica, R. nervulosa, R. exiguella (Plate
6, fig. 4), R. peruviana, R. reducta, and R. unilateralis. Those taxa with a comparatively distinct
double-structured cortex are R. australiensis, R. canariensis, R. filicaulis, R. fimbriata, R.
glaucescens, R. inflata complex, R. litorea, R. subfraxinea complex (Plate 6, fig. 5), and R.
tenella.
(b) Variation in the anatomical structure within the thallus. It was noticed when serial sectioning
thalli that differences in anatomical structure sometimes occurred depending on the portion of
the branch cut. Two forms of variation were found. The first form of variation is (i) variation in
the alignment of the hyphae in the supportive tissue in basal sections compared to the hyphal
arrangement in the mid and apical sections. Basal sections of R. celastri subsp. ovalis show a
typical thin outer cortex merging with the longitudinally oriented hyphae in the supportive
tissue, as found in the anatomy of R. celastri subsp. celastri (Plate 6, fig. 7). Sections cut through
the middle of the branch and towards the apex showed only a wide band of randomly oriented
<< Plate5 SEM photographs showing spore shapes in the Australian Ramalinae. Fig. 1 R. exiguella spores.
Scale 10 mm = 4 wm (X 2400). Fig. 2 R. inflata subsp. inflata spores. Scale 10 mm = 4 wm (X 2400). Fig. 3
R. inflata subsp. perpusilla (inland taxon) spores. Scale 10 mm = 4 wm (x 2400). Fig. 4 R. inflata subsp.
perpusilla (coastal taxon) spores. Scale 10 mm = 4 wm (X 2400). Fig. 5 R. celastri subsp. celastri spores
within the ascus. Scale 10 mm = 8-3 wm (X 1163). Fig. 6 R. litorea spores. Scale 10 mm = 4 wm (x 2400).
130 G. NELL STEVENS
0.5 mm
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 of R. inflata subsp. perpusilla thallus showing
variation from Fig. A — the supportive tissue being partly discontinuous; D. Algal clusters on the upper
side only, as found in some thalli of the R. inflata group; E. Cross section of R. whinrayi showing thick
outer cortex and irregular arrangement of supportive tissue which extends into the medullary region,
with isolated rafts of supportive tissue also present.
hyphae forming a network (Plate 6, fig. 6). This anomaly may be explained by the number of
laminal apothecia that this taxon produces on both sides of the branch. The lack of supportive
tissue and the random arrangement of hyphae could be due to the repeated formation of
apothecial tissue from the middle of the branch to the apex. Brandt (1906) found that variations
occurred within a single thallus, and illustrated this with serial sections along the branch of
Ramalina curnowii.
The second form of variation is (ii) variation in cortical structure between the upper side of the
branch and the lower side. This was noticed by Brandt (1906) in the species R. populina. In
Australian material such variation was found to occur in some specimens of R. inflata subsp.
inflata and R. inflata subsp. perpusilla, where the inner cortex on the lower side is narrow,
continuous and of even thickness, without an algal layer, but on the upper side it is partly
discontinuous, of uneven thickness with algal clusters interspersed between the ridges (Fig. 3D).
This formation appears to be a modification of the normal ‘farinacea type’ structure, to enable
optimal functioning of the phycobiont under particular environmental conditions. It is not a
constant feature of these taxa, so cannot be used taxonomically.
Another taxon which has an atypical and complex anatomy (Fig. 3E) is R. whinrayi. The
cortex consists of a thick outer cortex and a thick, discontinuous sheath of supportive tissue
made up of bundles with ridges extending across the medulla 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
Plate6 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 wm (X 6770). Fig. 2 Section
from outer to inner cortex — R. exiguella. Scale 10 mm = 1 wm (X 10450). Fig. 3 Usnic acid? crystals in
outer cortex — R. peruviana. Scale 10 mm = 1-8 wm (X 5682). Fig. 4 Section from outer to inner cortex
showing narrow outer cortex. Scale 10 mm = 5 wm (xX 2125). Fig. 5 Section from ouer to inner cortex
showing broad outer cortex. Scale 10 mm = 1:2 wm (xX 5000). Fig. 6 L.S. of mid-section of branch of R.
celastri subsp. ovalis. Scale 1 mm = 10 wm (X 105). Fig. 7 L.S. of mid-section of branch of R. celastri
subsp. celastri. Scale 1 mm = 19 wm (x 53). (Photographs for Figs 6 & 7 by courtesy of A. Henssen.)
almost totally lacking, or occur in patches appressed to the inner cortex. Ramalina fissa produces
very little medullary hyphae, whereas in some R. inflata subsp. inflata specimens the central
cavity is criss-crossed by arachnoid hyphae to the same degree as R. glaucescens, a species
regarded as having a solid, compressed thallus.
132 G. NELL STEVENS
The sorediate species belonging to the R. farinacea group in Australia have a thick felty
medulla compared to the R. farinacea taxa in the northern hemisphere which have a loosely
compacted but continuous medullary layer. Other sorediate species, R. canariensis, R.
fimbriata, R. reducta, and R. unilateralis, reveal large areas of exposed medulla which produce
soralia. The most compacted medullary hyphae are found in R. whinrayi (Fig. 3E).
Chemistry
Usnic acid is found in the cortex of the Ramalinae in varying amounts but it is not used
taxonomically as a diagnostic property. Atranorin occurs only rarely and in trace amounts so it is
not used for taxonomic purposes either. It is the medullary substances produced by the
Ramalinae — the depsides and the depsidones — which are important.
(a) The orcinol depsides: These comprise both para- and meta-depsides. The 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 (both para-depsides) or by sekikaic
acid (a meta-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
meta-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’-O-methylnorhomosekikaic, and 4’-0-methylnorsekikaic acids.
Cryptochlorophaeic acid is another orcinol meta-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-O-methylcryoptochlorophaeic acids
(Chester & Elix, 1978).
(b) The B-orcinol depsidones: Only four B-orcinol depsidones have been observed in Austra-
lian Ramalinae. Salazinic acid is present in four species and may be accompanied by trace
amounts of protocetraric acid, but protocetraric acid does not occur as a major component in
any of the species. Norstictic acid occurs in one coastal species and psoromic acid is present in
one inland species.
The number of depsidones found in the Australian Ramalinae is less than the number found in
Ramalaina species in Europe or North America, where fumarprotocetraric acid, hypopro-
tocetraric acid, stictic acid, protocetraric acid, norstictic acid, psoromic acid, and salazinic acid
all occur.
Salazinic acid occurs in both sorediate and fertile Ramalina species which inhabit the
Australian coastline (viz. R. pacifica, R. tenella, and R. fissa). In R. pacifica and R. tenella trace
amounts of protocetraric acid are sometimes present accompanying the salazinic acid. The
occurrence of these two acids in the same thallus is probably due to incomplete enzymic
oxidation of the side chain methyl group in the conversion of protocetraric acid to salazinic acid
(Elix, in litt.).
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 (1969a) 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, 1969b) and the explanation of the mixed acids would infer
hybridization (Brodo, 1978). But the islands are small and inaccessible and such a survey was not
possible during this study.
The occurrence of replacement compounds was also observed in the R. inflata group, where
lichen populations which are morphologically indistinguishable may contain either divaricatic
acid or sekikaic acid as medullary substances. Here again such taxa have been regarded as
chemical strains within the species; a decision strengthened by the presence of trace amounts of
sekikaic acid in some thalli which contain divaricatic acid. A third replacement compound,
salazinic acid, also occurs in the R. inflata complex. Biogenetically this depsidone is quite remote
from the two depsides, yet thalli possessing this acid are remarkably similar to those with a
depside chemistry. However, the depsidone taxon has been retained as a separate species (R.
fissa), as it has a geographically different distribution pattern from the depside containing
taxa.
(b) Accessory compounds: Accessory type compounds are also present in the R. inflata
complex, occurring sporadically in addition to the constant constituents, but having no
correlation with any morphological or distributional variations (Elix, 1982). The presence of
nordivaricatic acid in some divaricatic acid taxa and the presence of consalazinic or scabrosin
derivatives in the salazinic taxon have been regarded as instances of accessory acids accompany-
ing the major constituent and thus requiring no taxonomic recognition. Similarly the accessory
acid connorstictic has been found in the norstictic acid taxon in the R. subfraxinea complex.
Another Ramalina species complex comprising taxa with chemical variants of the replace-
ment type and with accessory compounds is the tropical/subtropical R. farinacea complex. The
acids involved are divaricatic acid (+ accessory compound stenosporic acid) and the sekikaic
aggregate of acids and salazinic acid, but this group is further complicated by chemosyndromic
variation occurring within taxa which produce the sekikaic aggregate of acids.
(c) Chemosyndromic variation: A chemosyndrome is a group of biogenetically related metab-
olites where one or two compounds are regularly the major components, and the minor
134 G. NELL STEVENS
biosequentially related constituent of one taxon becomes the major constituent of the other
(Culberson & Culberson, 1977; Elix, 1982).
In the R. farinacea complex, the two tropical/subtropical taxa R. nervulosa var. luciae and R.
nervulosa var. dumeticola contain the same acid components but these occur in different
quantities in each taxon (Stevens, 19834).
Correlation between secondary-product chemistry and ecogeography in the Ramalina
subfraxinea complex
The importance of chemistry in identifying taxa which have different amplitudes of ecological
tolerance has been stressed by Culberson (1967, 1969b), Culberson & Culberson (1967), and
Sheard (1978). The notable example of correlation between secondary-product chemistry and
ecogeography is found in the R. siliquosa species complex of the northern hemisphere. This
complex contains six sibling species each with a different depsidone chemistry (norstictic acid,
salazinic acid, stictic acid, protocetraric acid, hypoprotocetraric acid, and acid deficient).
Culberson (1969b) maintained that each chemical race was a different species which reflected
the physiological differences between the taxa. However, the R. siliquosa chemistry comprises a
replacement series of medullary depsidones which are able to be ranked by increasing numbers
of oxidation steps in their biosynthesis (Culberson, Culberson & Johnson, 1977).
The maritime R. subfraxinea complex in the southern hemisphere is similar to the maritime R.
siliquosa complex in the northern hemisphere as it is made up of taxa which produce six different
acids; four of the acids are depsides and two are depsidones (viz. sekikaic acid, divaricatic acid,
cryptochlorophaeic acid, boninic acid, norstictic, and salazinic acids), but these acids are not
closely related biosequentially (Elix, in litt.).
Morphologically only the salazinic acid taxon is distinguishable; the other taxa show only
slight morphological variations from one another, none of which is sufficiently unique and
consistent that the chemical constituents of every individual can be infallibly predicted from
appearance alone.
Culberson, Culberson & Johnson (1977) found a correlation between the chemistry in the
R. siliquosa taxa and different amplitudes of ecological tolerance of each taxon. Accordingly,
a study of the thalli belonging to the R. subfraxinea complex was undertaken in order to find
if a particular chemotype could be related to a particular type of habitat based on the
amount of exposure to the sea (e.g. exposed = on trees or rocks facing the open sea; sheltered
= on trees growing on the landward fringe of a mangrove community or on trees in clay pan
habitats).
In Table 3 the type of habitat (exposed or sheltered) at each location site is compared with the
percentage of thalli containing each acid found at the site. The table shows that there is a
difference in the percentage of certain acids at each site. The cryptochlorophaeic acid taxon
occurs in areas experiencing the greatest amount of exposure to the open sea, the boninic acid
taxon is generally found in exposed habitats (usually adjacent to quiet water), whereas the thalli
containing sekikaic acid, divaricatic acid, or norstictic acid are mainly found in sheltered well
protected areas away from salt spray. Salazinic acid containing thalli appear to tolerate a broad
spectrum of habitats from exposed to sheltered.
From Table 3 it can be seen that there is a definite tendency for different chemotypes to
occupy some microhabitats more commonly than others.
This environmental selection of different chemotypes into separate habitats is also apparent
to a larger extent along the more tropical coastline of Australia (e.g. north of latitude 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,
thalli collected from Lizard I. all contained salazinic acid.
Although the collections from these northern regions may not have been as numerous as those
to the south, enough material was collected at each site to make these findings plausible.
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.
2
o
e = =
2) ) ) ae
o & = = ) 3 <2 5 a a)
Z & 6 s 3 = 8 = ae ae
S — [=| — hd par) _— Le Be) oO
= S d. ey ele ey. Bien ee Kees
a 5 e Be Onna Bee ere oe ae
16 Daintree River sheltered —_ — 23 37 40 87
16 Mossman sheltered — 29 63 — 8 — 38
17 Mission Beach exposed — a> 10 — 15 — 20
18 Murray River exposed — 40 — 33 Ps | i
18 Hinchbrook I. sheltered 1 42 24 15: 16 2 82
19 Townsville exposed — a9 Pan, 12 — 7,
20 Bowen exposed 8 94 — 3 — — 53
Zi Hallidays Bay A. sheltered — — 32 68 — — 34
eA Hallidays Bay B. exposed — 94 — 6 — — an
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 Zo — — 60
24 Turkey sheltered 4 48 48 — — — 33
25 Burnett Hds exposed 10 90 — — — — 23
26 Hervey Bay exposed 36 64 — — — — an
5s Moreton Bay exposed 1 99 — — — — 100
29 Clarence R. exposed 66 34 — —_— — — 23
Distribution patterns in relation to climatic indices
The distinct distribution patterns of the Ramalinae indicate that there are differences in the
ecological requirements of each taxon. The composite map of the distribution of all of the taxa
(Fig. 1) shows that geographically the distributions lie in either the tropical or the temperate
zones with some taxa overlapping in the intermediate zones (e.g. subtropical and warm
temperate). In the higher plants it has been recognized that plants growing under tropical
conditions have different growth responses compared to plants growing under temperate
conditions, and that such responses are closely related to thermal optima for photosynthesis
(Specht, 1981c; Nix, 1982).
Asa knowledge of the major climatic indices which regulate plant response (e.g. temperature,
precipitation, and solar radiation) has proved valuable in understanding the distribution of the
phanerogram flora of Australia, it seemed reasonable to examine their value in understanding
the distribution of the Ramalina taxa throughout Australia.
Nix (1982) proposed that Australian plants can be classified in terms of their photosynthetic
response to temperature. His three major groups are relevant to the pattern of Ramalinae
distribution. (1) Megatherm Group — plants with photosynthetic optimum 26—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 multifactor Growth Index (G.I.). The Growth Index is different for plants with different
thermal growth responses. Figs 4A-B and 5A 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 tropical
Megatherm/Mesotherm
interzone 20-—24°C subtropical
Mesotherm 14—20°C
Mesotherm/Microtherm warm temperate
interzone 12-14°C
cool temperate
Microtherm <1
<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, SB).
(1) The megatherm element
Nine of the Ramalinae can be regarded as confined to a region with a megatherm Growth Index
value >0-30 <0-45. Their southern limits rarely extend below the tropics (latitudes 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 asa
mesotherm element because of its world distribution pattern which indicates its overall range is
in the subtropics (Stevens, 1983).
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
SS Growth index values > 0.45
E53 Growth index values > 0.30 < 0.45
(After Nix, 1981)
DISTRIBUTION PATTERN OF MESOTHERM PHANEROGAMS
TI) Growth index values > 0.45
anit] Growth index values > 0.30 < 0.45
(After Nix, 1881)
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.
G. NELL STEVENS
138
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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
coastlines.
In the south-west corner of Western Australia both divaricatic acid and sekikaic acid taxa
occur, reflecting the general aridity of this part of Australia even close to the coast.
There is a continuation of both mesotherm elements into Tasmania (Fig. 4B) and both R.
inflata subsp. inflata and R. inflata subsp. australis occur, but in different regions. The R. inflata
subsp. inflata taxa occur in the inland montane region whereas R. inflata subsp. australis taxa are
found in the drier eastern areas (Figs 21, 22). Sekikaic acid taxa do not occur in Tasmania
(extreme dryness is not a feature of the island). A microtherm influence could be responsible for
the distribution pattern of the R. inflata subspecies in Tasmania and in many instances the
presence of intermediate morphs causes difficulty in delimiting Tasmanian taxa.
Another taxon placed in the mesothermic group is R. celastri subsp. celastri. The majority of
collection sites for this taxon lie within the belt with G.I. values >0-45, with a few collection
localities to the west in the area covered by the mesotherm belt with G.I. >0-30 <0-45.
Specimens from the latter area are always broader than coastal specimens. This taxon extends
south to latitude 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
Australia (Fig. 20). It is also found in northern Tasmania.
Ramalina australiensis occurs coastally in the mesotherm pattern with G.I. value >0-45 (e.g.
Toogoom in Queensland and Yamba to Twofold Bay in New South Wales, latitudes 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 climate
region. Such environments are found in South Africa, Chile, and the Mediterranean (Raven,
1973) — all areas where this taxon occurs. The presence of R. canariensis at several sites along the
south-east coastline of Australia may be due to suitable microhabitat conditions.
The two species found on the Bass Strait islands (viz. R. whinrayi and R. caespitella) cannot be
ascribed accurately to any of the three response groups.
(3) The microtherm element
There are four taxa which occur in the mountainous areas of eastern and/or southeastern
Australia at altitudes exceeding 600 m which can be regarded as belonging to the microtherm
response group: (1) R. reducta, (2) R. glaucescens, (3) R. unilateralis, and (4) R. fimbriata.
(1) Ramalina reducta occurs at elevations over 1000 m in open-forests on the New England
tableland. The area experiences very low winter temperatures with snow. Figs SA—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
QV
Microtherm
GB CI> 0.6
EZZ7ACIt> 10045046
EE9 CI> 0.2<0.4 ee
E=a6r 2-6-2 E
Cs
BZ
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) Ramalina fimbriata 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 & @sthagen,
1980), East Africa (Krog & Swinscow, 1974, 1975, 1976), Fennoscandia and the British Isles
(Krog & James, 1977), Europe (Poelt, 1969), and the West Indies (Landrén, 1972).
Although the Ramalinae of ‘North America have not been completely revised, many of the
taxa have been recently investigated by Hale (1978), Rundel (1978a), Bowler (1977), Bowler &
Rundel (1972a, 1972b, 1973, 1974, 1978), Rundel & Bowler (1974, 1976), Hale & Culberson
(1970), and Moore (1968). Less recent studies cover the Hawaiian lichens (Magnusson &
Zahlbruckner, 1945; Magnusson, 1956); the Chinese Ramalinae (Zahlbruckner, 1930) and the
Japanese Ramalinae (Asahina, 1938, 1939).
The South American Ramalinae are in need of revision but records of taxa collected in various
regions of that continent appear in publications of Vainio (1890), Malme (1934), Follmann
RAMALINA IN AUSTRALIA 141
Ci ‘3
3 NG
Fig. 5B. Four Ramalina taxa with distribution patterns within the microtherm boundaries: 1 = R. reducta,
2 = R. glaucescens (sekikaic acid race), 3 = R. unilateralis, 4 = R. fimbriata.
(1967), Rundel (19785), Osorio (1970a, 1970b, 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
Table6 The global distribution of the Ramalina taxa occurring in Australia.
Islands of the Pacific
Ocean
Islands of the Indian
Australia
Africa
South America
New Zealand
India/Sri Lanka
Ocean
Tristan da Cunha
Europe
North America
West Indies
China
Ramalina species
*
australiensis
caespitella
canariensis
exiguella
filicaulis
fimbriata
glaucescens
litorea
peruviana
reducta
tenella
unilateralis
whinrayi
celastri
subsp. celastri
subsp. ovalis
inflata
subsp. inflata
subsp. perpusilla
subsp. australis
R. fissa
R. nervulosa
var. nervulosa
var. luciae
var. dumeticola
R. pacifica
R. subfraxinea
var. subfraxinea
var. leiodea
var. confirmata
var. norstictica
R. tropica
*** *£© &©* © & & & & He HR F
* & ©
*
*
ADDDAAAAAAAAARHA
*
*
*
*
*
*
*
*
*
* & & & * * * *
*
*
*
*
* & & &
*
*
* &* & & &
*
*
TOTALS 28 14 8 9 4 9 3 3 1 3 3 i
Antarctica to Australia was possible at this time (Table 7). Even in the late Cretaceous,
Australia still had connections with South America through Antarctica, but the migration of
plants via the southern route probably became progressively reduced until finally stopped by the
break of Australia from Antarctica in the Eocene (53-38 m.y. B.P.). The early break-away of
India from the rest of Gondwanaland (125-100 m.y. B.P.), and its rafting north across the
equator with the loss of some Gondwanaland species, could account for the lack of taxa similar
to the Australian Ramalinae.
Two major pathways of dispersal of the higher plants from west Gondwanaland have been
postulated by plant biogeographers: (1) a temperate migration route via Antarctica from west
Gondwanaland (i.e. South America and Africa), to Australia and New Zealand, and (2) a
tropical route from East Africa to Madagascar, the islands of the Indian Ocean and Indo-
Malaysia to Australia (Croizat, 1952; Raven & Axelrod, 1974). As the Ramalina species found
in Australia can be placed in a range of temperature response groups from cool temperate to
144 G. NELL STEVENS
Table 7 The history of the break-up of Gondwanaland, indicating available migration routes (from
Raven & Axelrod, 1974 and Powell, Johnson & Veevers, 1981).
Million
years BP Geological periods Event/geographic situation
125 early Cretaceous Africa and South America begin to split; India begins to split
from Australia/Antarctica but remains attached to
Madagascar.
110 + 10 Direct migration between west Gondwanaland and Australia
ceases.
100 mid Cretaceous 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.
90 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.
80 late Cretaceous Australia/Antarctica extend from 40°S to the polar region.
South America closely connected with Antarctica 80—30
million years BP.
Separation of New Zealand and New Caledonia from
Australia.
63 early Palaeocene Africa and Europe connected at Spain and possibly with Asia
at Arabia, allowing both northward and southward
migration.
55 Palaeocene-Eocene Oceanic crusts form between Antarctica and Australia;
northern edge of Australia now at 30°S and commencement
of separation of Australia from Antarctica.
49 mid Eocene 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.
45 Direct migration between India and Asia.
30 mid Oligocene The leading edge of Australia as it rafts northwards still south
of the latitude of the Sunda Arcs.
20 early Miocene Initial contact between Sunda Arc and New Guinea
(abundant precipitation).
10 late Miocene Collision of Australia with the Sunda Arcs.
Antarctic ice-sheet rapid expansion (lowered precipitation).
Present Holocene 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 Jorgensen (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 (108-100 m.y. B.P.). Fossil evidence indicates that many of the Gondwanan
families and genera had occupied Australia by the early Tertiary (63-55 m.y. B.P.) (Specht,
1981c).
(a) Step by step dispersal: Schuster (1979) stated that migration of land plants normally tends to
be by short range dissemination as members of structured communities, and he proposed that
the dispersal of Hepaticae occurred as part of the dispersal of whole plant communities. It is also
reasonable to hypothesise that the Ramalina species also made this step by step spread across the
continents of Laurasia and Gondwanaland with the migrating structural communities during the
Cretaceous-early Tertiary.
This suggested step by step migration of plant communities appears to be a valid means of
dispersal as this enables plants which have similar habitat requirements to expand as a whole, in
favourable climates, and so migrate when conditions are favourable and disappear from areas
where environmental conditions have deteriorated. It seems highly likely that under the
circumstances lichens which grow within these communities would migrate at the same rate and
in the same direction.
If step by step migration halted with sea-floor spreading of the continents, then another means
of dispersal has to be proposed. Seeds of the higher plants have been found to be carried by birds
and animals over long distances; the lighter seeds being wind dispersed. The Ramalinae produce
both spores and vegetative diaspores and it is feasible that these could be wind dispersed or
carried by birds.
(b) Wind dispersal: If long distance dispersal of spores by the wind is accepted then this means
of dispersion would have been in operation in early times. It is reported by Raven & Axelrod
(1974) that with the separation of Australia and Antarctica in the Eocene, there was a
146 G. NELL STEVENS
strengthening of the circum-Antarctic wind which could have carried spores from South
America to Australia, Tasmania, and New Zealand.
The success of a spore in a new environment would need the availability of a suitable
phycobiont and the lack of competition from plants already in the area. With the vegetative
diaspore both partners arrive together so that provided the environment is hospitable to their
genetic requirements they will readily become established in the new habitat. Spores can
become established after long distance wind dispersal since fertile lichen species have been
found on the islands of Hawaii. Jorgensen (1979, 1983) remarked on the efficiency of wind
dispersal of lichen spores in the population of the remote island of Tristan da Cunha.
If the original dispersal of the Ramalinae is comparable with the original dispersal of
angiosperms and gymnosperms, then their migration from west Gondwanaland may have taken
place by both step by step migration and wind dispersal, with the latter means continuing after
the break up of the continents.
(4) Centres of origin
The centre of origin of a group is often regarded as that area having the greatest concentration of
species. Hale used this method in assessing the centre of origin of the genera Relicina (Hale,
1975), Pseudoparmelia (Hale, 1976a), Parmelina (Hale, 1976b), and Bulbothrix (Hale, 1976c).
If the numbers of Ramalina taxa occurring in each landmass that once formed Gondwanaland
are estimated (and these numbers must be only approximate as without recent revisions many
names could be synonyms or misidentifications) then it is found that 41 Ramalinae occur in
South America, 42 in Africa, 36 in India, and 30 in Australasia.
South America and Africa (once West Gondwanaland) appear to have the largest number of
Ramalinae and could be regarded as the centre of origin of the southern hemisphere taxa.
The occurrence of high numbers of taxa within the one genus, in a particular region, can,
however be due to a re-radiation of species after the original dispersal of the taxa. When an
environment is free of competitive species, newly evolved taxa belonging to the genus will
become established. This situation has been found in Western Australia amongst the eucalypts,
where secondary radiation of species has occurred in a region where there is little competition
from other genera (Pryor, 1981).
Such a situation may have arisen in Australia with the evolution of the mangrove flora in this
region (Specht, 1981b). Mangroves are the major phorophytes for the present day R. subfrax-
inea complex. It is proposed that the ancient stock of the R. subfraxinea group could have
entered Australia via the northern route from East Africa (as this group is regarded as a
megatherm response group) and become established along the Australian tropical coastline on
the newly evolved mangroves. The occurrence of all six chemotypes in the region around
latitude 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 Australia R. inflata subsp. inflata morphology closely resembles some
forms of the East African taxon R. calcarata, but their chemistry differs in the minor
constituents.
RAMALINA IN AUSTRALIA 147
Chemical difference is the only character which separates the Australian taxon R. filicaulis
from the American taxon R. anceps. Material similar to the Australian material, and also lacking
medullary acids, has been cited from South Africa. It is proposed that all three continents
possess species which could be regarded as vicariant taxa belonging to the R. usnea complex, as
they are ecologically and morphologically similar.
On the basis of the evidence discussed above it is postulated that Australia has been populated
by species of Ramalina since the late Cretaceous. Since west Gondwanaland is accepted as the
centre of origin for the angiosperms, based on fossil evidence, there is the possibility of these
acting as phorophytes for ancestral taxa of the present day Ramalina, as that same region is a
likely centre of origin for Ramalina. The step by step spread of the plant communities would
have enabled the Ramalinae to spread by one of two migration routes either (1) the southern
route from South America (after Africa had separated) via the Scotia arc, Antarctica to
Tasmania and Australia or (2) the northern migration route from East Africa via Madagascar,
the Mascarenes, Indo-Malaya to northern Australia, by which means the megathermic Ramali-
nae could have reached tropical Australia. Once established in Australia the megatherm
Ramalina taxa remained a maritime group as inland habitats would have been inhospitable.
Secondary radiation from Australia by some of the megathermic taxa appears to have occurred
with the spread of species to the Pacific and Indian ocean islands.
The microtherm Ramalinae became established in the mountainous region of southern
Australia since these habits would have suited their thermal response patterns as Australia
drifted northwards and the temperatures increased.
The climatic changes that occurred in the late Miocene Period caused the desiccation of the
continent. Subsequent speciation of mesotherm Ramalinae could have occurred producing taxa
which occupied the new and dryer habitats of the south-west and west of Australia.
In this way the present distribution pattern of the Australian Ramalinae reflects their
palaeobiogeographic and palaeoecological history.
Taxonomy
The delimitation of Ramalina species proved difficult owing to the amount of morphological
variation shown by each taxon. A large amount of material had to be examined in order to
ascertain what concepts should be applied in the delimitation of each taxon. Three taxonomic
ranks have been used.
Taxonomic ranks
(1) Species: The rank of species has been given only to those taxa which can be recognized by
several distinct morphological traits which collectively produce a particular thallus type (allow-
ing for an appropriate amount of variation), together with a set of specific anatomical and
chemical properties, and which show a distinct habitat and distribution range.
Difficulties arose when two or more chemotypes produced a single morphotype; this problem
had to be resolved by considering the habitat and distribution range of the taxa, and if the only
difference found was in the chemistry then these chemical races were treated as conspecific.
The concept of species pairs (Artenpaare), as proposed by Poelt (1970, 1972) is that species
which reproduce by vegetative methods (secondary species) have been derived from existing or
extinct fertile species (primary species), and that primary and secondary species have identical
(or closely related) chemical components. This view is held by most lichenologists and species
pairs have been found in many genera e.g. in the Physcia and allied genera (Moberg, 1977),
Parmelia sect. Hypotrachyna (Culberson & Hale, 1973), Dirina (Tehler, 1983), and to a lesser
extent in Alectoria (Brodo & Hawksworth, 1977).
Only a few records of species pairs have been found in the Ramalinae. Rundel & Bowler
(1976) discussed R. leptocarpha and R. sublectocarpha as a fertile-sorediate species pair and
Krog & @sthagen (1980) have commented that the fertile species R. implectens may be regarded
as the parent morph of the sorediate species R. farinacea.
148 G. NELL STEVENS
None of the Australian sorediate Ramalinae is regarded as derived from any of the fertile taxa
found in Australia today.
(ii) Subspecies: The delimitation of a taxon based on well marked discontinuities in characters
was not always possible. Some Ramalina taxa were found to intergrade into each other, differing
only subtly morphologically yet extending over large areas. When such complexes were
examined carefully the most obvious morphological changes were found to coincide with
different geographical regions.
The term subspecies has therefore been applied to a taxon which could be shown to occupy a
particular region within the total distribution range of a large complex (as defined by Hawks-
worth, 1976). This taxon can be extremely variable within its geographical confines and can
produce several ecotypes which occupy separate habitats.
As subspecies intergrade into each other, their delimitation is much more arbitrary than that
of a species, and depends more on geographical divisions than on chemical or morphological
differences. Imshaug & Brodo (1966) pointed out that the use of the rank of subspecies
permitted a classification which is not only practical but one which indicates a possible
phylogenetic system. The evolutionary relationships that may exist in some of the Australian
taxa have been indicated by the use of the rank of subspecies.
(iii) Variety: The rank of variety has been used when more or less morphologically identical
taxa have been found to produce different chemistries which correlate with distinctive distri-
bution patterns.
The production of either divaricatic acid or sekikaic acid in populations of the one taxon posed
a nomenclatural problem. Several lichenologists have mentioned the same difficulty e.g. in the
genus Dirinaria (Jorgensen, 1974) and in the genus Ramalina (Imshaug, 1972; Rundel, 1978a).
The solution has been to regard both acids as representing chemical races within the one taxon.
Hybridization
It is not easy to establish the genetic homogeneity or heterogenity of lichenized fungi. Jahns
(1974) pointed out that thallus or hyphal fusions do occur in lichens, which could enable a form
of hybridization to take place. A specimen of R. leiodea recorded by Elix (1982) as being a
‘schizophrenic thallus’, had ‘left-hand branches containing one acid, the right-hand branches
another and the central branches containing a mixture of both’. In this case the two acids
involved were two depsides, not closely related (cryptochlorophaeic acid and boninic acid).
Another such combination of two acids in one specimen belonging to the R. subfraxinea
complex has been found during the present study; the specimen was collected at Hinchinbrook I.
where all acid races of the R. subfraxinea complex are present. The thallus concerned contained
both boninic acid and salazinic acid, one a depside and the other a depsidone. As these two acids
are not biosequential metabolites it appears likely that this is a similar form of hybridization to
that found by Elix (1982), and that both could be regarded as ‘chimeras’ produced by the fusion
of hyphae from two different spore types. It is not known what chemistry the thalli produced
from spores of this plant would have. No other thalli with this combination of acids has been
found.
The occurrence of sekikaic acid in the divaricatic acid race of R. subfraxinea var. subfraxinea is
regarded as different from the above. Sekikaic acid and divaricatic acid are reasonably closely
related biosequentially and apparently can interconvert (Elix in litt.) which would result in
chemical combinants if the conversion was incomplete. However the hypothesis of interspecific
hybridization (Brodo, 1978) remains feasible and has to be considered beside the hypothesis that
incomplete conversion of a precursor acid occurs during the biosynthetic steps from one acid to
another (Bowler & Rundel, 1978). If hybridization is accepted as the cause of this mixture of
acids then it would be natural to expect the proportions of each acid to be equal in the thallus.
However, this was not the case with the taxa examined; in all instances divaricatic acid occurred
in far larger quantities than did the sekikaic acid. It is therefore suggested that incomplete
conversion of one acid to another has taken place.
RAMALINA IN AUSTRALIA 149
Table 8 Taxonomically important characters in the Australian Ramalina taxa.
major acid constituents
orcinol depsides B-orcinol
depsidones
substrate thallus para- meta-
cryptochlorophaeic
saxicolous
inflated
sorediate
divaricatic
evernic
sekikaic
boninic
salazinic
norstictic
psoromic
Ramalina species
>| nil acids
. australiensis
. caespitella
. canariensis
. exiguella
. filicaulis
fimbriata
glaucescens
litorea
. peruviana
. reducta
. tenella
. unilateralis
whinrayi
Species complexes
R. celastri
subsp. celastri
subsp. ovalis
R. inflata
subsp. inflata
subsp. perpusilla
subsp. australis
R. fissa
R. nervulosa
var. nervulosa
var. luciae
var. dumeticola
R. pacifica
R. subfraxinea
var. subfraxinea
var. leiodea
var. confirmata
var. norstictica
R. tropica
mx
*
~
mx
*
xx > > +! Corticolous
Pe
~<
~~ X
D4 4 D4 Od DS OS DS DS DS DS DX | Solid
~ mK XM
DDD DW WD RWDWD WDA
~*~ x ~*~ x
mK
mK mK
»* mK x
~*~ ~
mK xX
~ KK
~ X
Xx
mmm MM KKM
Mmm MM
*
*
X
Description of the genus Ramalina
Ramalina Ach., Lichenogr. Univ.: 122,598 (1810), nom. cons.
Thallus fruticose, greenish-grey, pale green or greenish-yellow, caespitose, erect, subpendulous
to pendulous; branching dichotomous, subdichotomous to irregular or dense intricate; branches
few to many, twig-like, strap-like to palmate, flat, subterete or terete, solid or inflated; apices
forked, attenuate, rounded; surface shiny or matt, smooth or rugose, pseudocyphellate;
holdfast delimited or dispersed; soralia small and punctiform to large and round or elongate;
150 G. NELL STEVENS
cortex composed of two layers, the outer cortex hyphae more or less anticlinal or randomly
oriented, the inner cortex hyphae periclinally arranged; hyphae surrounded by a large amount
of matrix; medullary hyphae loosely interwoven between the upper and lower cortex or confined
to the algal zone.
Apothecia terminal, subterminal, marginal, lateral, laminal; convex, plane or concave,
sometimes incised; disc pale to flesh coloured; margin concolourous with thallus; asci clavate,
containing 8 spores; spores hyaline, oblong, ellipsoid or fusiform, straight or curved, 1-septate.
Pycnidia inconspicuous, pale, usually rare in Australian species.
The algae are Trebouxioid, globose, forming an irregular ring along the edge of the inner
cortex; when confined to the upper side only the thallus becomes dichroic with a white lower
surface.
Type species (conserved): Ramalina fraxinea (L.) Ach.
Enumeration of the Australian taxa
The Australian Ramalinae comprise 28 taxa which can be grouped into 17 fertile and 11
sorediate taxa. Sixteen species have no subspecific taxa recognized, and four species contain a
total of 12 subspecific taxa.
Table 8 lists the taxonomically important attributes of each of the 28 taxa; those belonging to
species complexes are placed after the alphabetically arranged single species, as is also done in
the description of the taxa. The disposition of the other Ramalina taxa recorded by Wetmore
(1963) and Weber & Wetmore (1972) is shown in Appendix 1.
The specimens examined are cited with location, latitude and longitude in degrees, type of
substrate, date, the collector’s name and the herbarium abbreviation (Holmgren, Keuken &
Schofield, 1981). Collectors initials have been used for those collectors whose specimens are
cited most frequently in this work. These names have been abbreviated as follows:
AA A. Archer JC J. Cashin
ED E. Dahl JW J. S. Whinray
GB GoC. Bratt NCS N. C. Stevens
GK G. Kantvilas NS N. Sammy
GNS G. N. Stevens RF R. B. Filson
HS H. Streimann RS R. D. Seppelt
JAE J. A. Elx RWR R. W. Rogers
Only one specimen is cited from each location for most of the taxa; where the taxon grows
profusely over a wide area, only specimens from selected sites are listed: viz, R. celastri subsp.
celastri, R. celastri subsp. ovalis, R. inflata subsp. inflata, R. inflata subsp. perpusilla, R. inflata
subsp. australis, and R. fissa.
Key to Ramalina in Australia
la. Thallusinflated (partially or totally). c0cc5 vse sees step castes 0 ce cn water ieecayenesaoeseee recone 35
ib: Uhalhws mori ated Saka sthek eee es es coe Ree Pe pentcr ties sue oct tee a nan cea nao 2
De (1b): Plant SaxicOlous. os. s-c052. so serccaes so Aipocsecae ccs san ssnecusanae Ropes bene austadnaeenr te duekoads Beeeds 3
2b. PIANECONTICOIOUS «29 oecnetwteciev se tate een tadaccutis des sncuncccoauchrenvnrwerotanereeriets 12
Sa (Zaye Dhalltis SOTECIAtE co5.o coe os seas fens eon sade teks fe emicibei aa pe alot eren ae tise SACRA EMO ONe OU: yamoe nated 4
3b. Ralls MOCSOLECIALG A ecco nse teen oe ot tata ons oe ce nee ach eetigecinis tun oseeeae pe anginne? 5
4a (3a). Soredia eruption through eroded lower surface and at apices ......... 6. R. fimbriata (p. 161)
4b. Soredia contained in well defined soralia, which are marginal and sometimes laminal
2. R. caespitella (p. 155)
5a, (50). Phallus made up Ola few 1OO8e DIGNCHES. 5. ¥,..cs.tacacosscesnery oxioes eventos orenacunadanateczee sts 6
Sb. Thallus composed of numerous branches forming a dense cushion ................0c:0eeeees 8
6a (5a). Branches compressed, flat (apothecia marginal) ...... 14. R. celastri subsp. celastri (p. 180)
6b. IBTABCNES SUDTCTOLE TO LOLOLS fociccdsd.sca cs noinisen stboiaspwansts aesgiseriaees sss enesenrpernentnntsesa i
7a (6b). Branching dense, distal, K— (no medullary acids) .................. 1. R. australiensis (p. 152)
7b. Branching sparse, K+ red (salazinic acid) ...............sscsesessceseseeees 20. R. tropica (p. 210)
8a (5b).
8b.
9a (8a).
9b.
10a (8b).
10b.
11a (10a).
11b.
12a (2b).
12b.
13a (12a).
13b.
14a (13b).
14b.
15a (14b).
15b.
16a (15b).
16b.
17a (16a).
17b.
18a (16b).
18b.
19a (18b).
19b.
20a (19a).
20b.
21a (12b).
21b.
22a (21a).
22b.
23a (21b).
23b.
24a (23a).
24b.
25a (24a).
25b.
26a (25b).
26b.
27a (26b).
210;
28a (27b).
RAMALINA IN AUSTRALIA
ove Ti oly Verh <6 (8 ONRRp re RES Nee Papen Be Sys anne EE oy Ne a oR Mca RiP ARRON Ey Beh Ae
Branches inflated, Omen Sout 2200.74.55 cis pass- eos cu se oey fees sacsseeie eae oncineoncod tavanvnti eg ans
Apothecia terminal, large 3—10 mm diam. ..............ccssoeesecteceens
Apothecia subterminal (subtending narrow pointed branchlets, apothecia 1-2 mm
diam.)
Thallus showing some inflation; splitting and tattering of branches common ..............
Teak US FOE WEL ACEO cas 2 hs cic ioe es dees atc kd Sans sean Nnnmenec ees un euguae aaiie Lecenedoae
WMedulla K+ £60 (SAlAziOtG RCIA) ian ee a vese so sece cs cen sg heer
Medulla K— or K+ pink (divaricatic acid or sekikaic acid)
15. R. inflata subsp. australis (p.
SH ALTSISOTECIALC eee iae ee career cae ae aoa eee Te EE aes, eerie eats enone
PNAS MOCSOTECIALC cee eit a eee ae rere oie a ch aI te eee tata toes Lamemare
Lower surface of anastomosing strands resting on the medulla........
Lower surface not of anastomosing strands Only... 7c255. 0 arise .hd scien sa vensevccevnsdee seasons
Branches few, usually broad at base (soft textured, soralia apical and marginal)
3. R. canariensis (p.
Branches numerous, narrow
Branches usually cartilaginous (splitting open at intervals along their length)
12. R. unilateralis (p.
Branches not cartilasinous, Completes: co.cc sh sscnes Goenan ave ue eeaonsninda sabesoees aabeds
Meédulla K+; red (salazinie acid). 3c aeewa visa sewed oiesace tee ges Dione pnetnee eens ces oa a
Medulla K— or K+ pink
Soralia punctiform, mainly apical on branchlets (thallus up to 3 cm high)
11. R. tenella (p.
Soralia ellipsoidal or round, marginal on elongate branches (thallus usually 4-8 cm
long)
Soralia ellipsoidal or round, marginal on elongate branches ...................00ccecseeeeee ees
Soralia punctiform, marginal and apical (branching dense, intricate, thallus resilient in
texture) 9. R. peruviana (p.
Medulla K+ pink (sekikaic aggregate acids)
Medulla K— (divaricatic acid and stenosporic acid) 17. R. nervulosa var. nervulosa (p.
Sekikaic aggregate acids + faint ramalinolic acid
Sekikaic aggregate acids + faint homosekikaic acid ....
Seem mee meee eee meee meee eee e eee eee eeeeeeeeseeeeeeeesseesssessssEssssessesseee
Branches flat, sometimes canaliculate, containing norstictic acid (spores 12-16 x 4—5
MATE Sale Parse erica cies oan ote Ove anon ees
Branches usually subterete, never canaliculate, containing salazinic acid (spores 14-22
Me BN MON) oy sascitos atevex sa vestadddiere Gee Miswies ra ere eed ack tale seeae as
Branches subterete to terete, containing no medullary acids ..................:ccceceeeeee eens
Branches fiat, with'or without medullary acids .22..<..ccscccssancrenvansiwatencteoateasnareaneade
PLANS Bo an vores ie tacercd oaalnn Gores combine viouaiek ieyiecusrnpnlaeicus ov owe ptaee Rac IRR AS Cedar e SORE aaa aYe
Medulla K+ pink
Apothecia lateral and subterminal with branch apex appearing as a long spur (some-
TINGS DISK MOVE): cers waco ere toes eyiniendicty ores ceeaasseeencsnntes
Apothecia never subterminal nor bearing a spur
Distal branchlets occur on otherwise unbranched primary branches
1. R. australiensis (p.
Dense distal branchlets not present -[oc6. sec ssecs nose cous thy sen cre edeccdaderanee heevers ene vises
Thallus thread-like (with dichotomous branching)
Thallus fruticose
Apothecia marginal on both edges (branches may be canaliculate, no medullary acids)
14. R. celastri subsp. celastri (p.
13. R. whinrayi (p.
8. R. litorea (p.
18. R. pacifica (p.
17. R. nervulosa var. dumeticola (p.
17. R. nervulosa var. luciae (p.
19. R. subfraxinea var. norstictica (p.
20. R. tropica (p.
4. R. exiguella (p.
152
G. NELL STEVENS
28b. Apothecia not matginal On DOUR CORES «<< tec ceassne sens va ndunvaet ones espe vespaaWagamas tenes ee 29
29a (28b). Apothecialaminal, numerous, and equal in size ......... 14. R. celastri subsp. ovalis (p. 184)
29b. Apothecia not laminal and numerous and equal in Size ...............eceecececeeseceeeeeeeseees 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
BPP OURC CUMIN oi sac wied th ars sates oceamisd cabenay<otadaccvevaorreg utnasetnen evens uate reeeaee eter 31
31a (30b). Branches narrow with dichotomous branching (apothecia marginal or subterminal or
in the axils of branches, small 0-5—1-5 mm diam.) .................. 7. R. glaucescens (p. 163)
31b. Branches narrow to broad, branching usually from the base only .....................0c00e0e 32
Bal SLD). MICGUUNA RF Pik: 22 .carusl suesdindaseceds tiaes tale aiaaus Seavsssaqans stcrccse see stk es pecereee tee nes 33
32b. INSOUT e ooanog oipe eo ssa sac se tango taaeabedaed sanasens Un sdt4d pee eee es dala den tee eeeeane nents 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 ......................066 19. R. subfraxinea var. leiodea (p. 207)
34b. Thallus containing divaricatic acid ................ 19. R. subfraxinea var. subfraxinea (p. 203)
Soar (lay: a mallleSOrediatens:. cores. seccee sik ne lea tes fcacos cena euoweteodassageeuuseeaes 6. R. fimbriata (p. 161)
35b. alls not Sorediate ccc sont eteaete cette oua cs fr aitea ce croet cee reac rege seueceseere cast tect yak 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. inflata subsp. australis (p. 191)
36b. Thallus not button-like nor bearing apothecia between sterile branches .................... af
STA SO)] ““DLAMCHES SONIUOPEN ANG Ca tlenee 5h runsiaacsaio cs nanlccaer ou ety ony sears evel ease an aaaoesuacetees 38
37b. Branches perlorate Out COMPIOLE SS sc nccssiey aancicers ane tumrse teenies ta sip wun egraddeuctanas cteeeee 39
38a (37a). Thallus grossly inflated, few or no perforations, splitting open of branches, medullary
ACIS GivaniCaticOrsekikaic.. 5. /.25500.1s<s00 caaeoresesees 15. R. inflata 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. inflata subsp. perpusilla (p. 188)
39b. Thallus totally inflated, apotheciaconcave andinnate 15. R. inflata subsp. inflata (p. 185)
1. Ramalina australiensis Ny].
Plate 7, fig. 1; Plate 13, fig. 1.
in Bull. Soc. Linn. Normandie I, 4: 120 (1870). Type: Australia, West Australia, Swan River, 1846,
Verreaux s.n. (PC! — holotype; H-NYL 37491! — isotype; usnic acid).
Ramalina myrioclada Mill. 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(—1-0) 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
Plate7 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
~
Ky, rey yrrrye
IUNUERUUUVUAUUUINUNNNUUUVUUNIONOONTONUORUONUOUEURTOVOVIMM il OTOL DOL DOU TOOOVUOTDU
¢
Ton TITITITLI LIL
154 G. NELL STEVENS
0-5—1-0(—1-2) mm diam., plane to convex, margin entire; spores ellipsoid to fusiform, straight,
12-16(—20) x 4-6 wm (Plate 4, fig. 2).
Chemistry. Usnic acid only.
Remarks. The name R. australiensis as the oldest name has been upheld for this taxon,
overriding the more commonly used name R. myrioclada. However the location given for the
type material - Swan River, Western Australia — is dubious. All recent collections have been
made in eastern Australia with no other material being found in the west. Whether the original
material from Western Australia was wrongly recorded was not able to be ascertained.
The type material of R. australiensis has branchlets with black tips but this feature (as with R.
exiguella) is not present in most specimens. The type is a coarse thallus form of this taxon, rather
resembling that of the subtropical Queensland collection. There is not sufficient type material to
obtain an overall picture of the total thallus.
The degree of secondary branching varies from specimen to specimen, those with dense
branching resemble R. peruviana whereas others have sparse branching producing an open
thallus form resembling the morphology of R. filicaulis N. Stevens.
The names R. usneoides and R. usnea have been applied to some specimens of R. australiensis
collected in New South Wales but the Australian taxon can be distinguished by its branching
pattern and subterete branches instead of the characteristic flattened branches of R. usnea
sens. str.
Distribution and habitat. This subtropical/warm temperate maritime species does not have a
120 130 140 150
10-
20-
a ote a TTS Ras
30-
40-
ee | rn L rt
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. australiensi: 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, JAE (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, 151°10’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. Saenger (BRIU2137NS).
2. Ramalina caespitella N. Stevens Plate. fig, .2.
in Lichenologist 18: 183 (1986). Type: Australia, Long Island (Bass Strait), 1969, Whinray s.n. (MEL! —
holotype; sekikaic acid, 4’-0-demethylsekikaic acid, + several terpenes, and usnic acid).
Thallus saxicolous, pale greenish-grey to stramineous, minute, caespitose, rigid, up to 1-0 cm
high; branching subdichotomous, irregular, narrow thalli densely branched, broad thalli
sparsely branched; branch width 0-1—2-0 mm, exceptions to 6 mm, branches solid, flat to
subterete, narrow branches nodular, apices broad and blunt; surface shiny; holdfast delimited or
diffuse, soralia marginal and laminal, round to ellipsoidal, becoming fissural on the upper
surface occurring on eroded areas of the lower surface and at the apices. Apothecia rare, laminal
near the branch apices, rarely terminal, disc 1-0 mm diam, concave, margin thick, entire; spores
not produced.
Chemistry: sekikaic acid (M), 4’-0-demethylsekikaic acid (t), + several terpenes and usnic acid.
Remarks. Ramalina caespitella produces two morphotypes which intergrade into one another.
One morph has narrow subterete to terete nodular branches with apical and marginal soralia,
whilst the other morph produces very thick, flat to subterete branches bearing numerous soralia
on both upper and lower surfaces. The extreme button-like morphology of both forms is
indicative of their exposed, saxicolous habitat. No other taxon has been found to have a close
affinity with R. caespitella.
Distribution and habitat. This temperate maritime taxon is found on several of the islands in
Bass Strait and along parts of the 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, JW (MEL1012702); Deal I, 39°30’S, 147°22’E, on
granite, 1972, JW (MEL1012940); Furneaux Group: West Sister I, 39°45’S, 147°56’E, on granite, 1966, JW
156 G. NELL STEVENS
t { 7 1
120 130 140 150
B aj 10
0 800km 8
20+
ye PS aed
AQ ¢
9
1)
30+
=
GES
a 407
uy \ L L
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, JW (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°05S’E, on rock, 1974, JW (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.
in Ost. bot. Z. 9: 8 (1904). Type: Gran Canaria, Tafira, Bornmiiller, Pl. Canar. 3500 (W! — isotype;
divaricatic acid and usnic acid).
Thallus corticolous, pale green to grey green, erect to subpendulous, 1—3(—5) cm long;
branching palmate or irregular; branch width 1-3(-10) mm, apices blunt, usually split with
margins separated; surface matt, smooth to rugose, sometimes coarsely reticulately ridged by
chondroid strands, pitted distally; holdfast delimited; soralia marginal and apical occurring
through the separation of the upper and lower surfaces.
Apothecia not seen. Reported as rare.
Chemistry. Divaricatic acid, usnic acid.
Remarks. Fine thalli of this taxon come close to resembling immature thalli of R. unilateralis
RAMALINA IN AUSTRALIA 157
and both species contain divaricatic acid, but the cartilaginous surface and the finely lacerated
apices of R. unilateralis enable the two to be separated.
Distribution and habitat. The distribution pattern of R. canariensis in Australia is mainly
maritime in areas experiencing a Mediterranean type of climate (Fig. 10). The only inland
collections are two South Australian locations (Springton and Kuitpo Forest) at elevations of
300 m where frequent mists occur.
Krog & @sthagen (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’. Insome
of the drier Australian coastal sites morning fogs are prevalent.
Along the southern coastline of Australia this species grows in well-lit situations protected
from winds, usually on twigs of Leucopogon parviflora, Casuarina, and Melaleuca spp. It has
also been found on mangroves (Avicennia marina) along the eastern coastline south from
latitude 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.
120 130 140 160
0 800km sy Uy
, m . 1 1 a)
&
f 20~
Qr
9
a
f 20,
S
v e
Cm
9 @
é
40N
1 T i ie
Fig. 10 Distribution of R. 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, 150°10’E on A vicennia marina, 1978, GNS (BRIU2171NS); Batehaven Beach, 35°45’S, 150°13’E,
on Casuarina sp., 1975, JAE (JAE 1061); Burrenwarra Pt., 35°49’S, 150°14’E, on Casuaria sp., 1975, JAE
(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, J. 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, on Melaleuca sp., 1970, RF (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, JW (MEL1025619); West Sister I, 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. Ramalina exiguella Stirton Plate 7, 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 (S5—)6—8 um (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 R. attenuata.
It could be that this group of taxa is a complex with several chemical races, but until the type
material of all of these species can be located and examined, the relationship between these taxa
cannot be resolved. Stirton stated in his type description that R. exiguella had affinities with R.
melanothrix Laurer, but the latter species is quite distinct and the type material does not
resemble Australian material of R. exiguella.
Distribution and habitat. The distribution of this coastal species is continuous along the eastern
shoreline of Australia from latitudes 23°S—35°S, with additional occurrences at 18°S and 21°S. 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, 21°06’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); Fraser I, Wathumba Ck, 24°50’S, 153°13’E, on Ceriops tagal, 1975, GNS
120 130 140 160
KG, 104
cede L 1 1 800km
G
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204
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Fig. 11 Distribution of R. 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 (BRIUS459RR); 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 (BRIU4332NS); 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 filicaulis N. 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
plano vel convexo, ad 0-5—1-0 mm in diametro; sporae ellipsoideae vel fusiformes, rectae vel raro
curvatae, 12—16(—20) x 4-6 wm. Acidum usnicum tantum continens.
Holotype: Australia, Queensland, Lamington Plateau, 9 km N of O’Reillys in rain-forest on Araucaria
cunninghamii, alt. 920 m. 27 November 1983, G. N. Stevens 3730 (MEL 1048085!).
Thallus corticolous, green-grey, pendulous or decumbent, up to 20 cm long; branching
predominantly dichotomous, curving at the axils; branches width 0-2—0-5 mm, narrow, thread-
like, subterete to terete, tapering, apices attenuate and filamentous, often hooked; surface
smooth, with linear pseudocyphellae; holdfast small, delimited but branches often attached to
the substrate along their length; soralia absent.
Apothecia uncommon, lateral sessile, disc 0-5—1-0 mm diam., plane to convex; margin entire;
spores ellipsoid to fusiform, straight or rarely curved, 12—16(—20) x 4-6 wm.
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). Landr6én (1972)
recorded R. anceps as a species in the Caribbean Islands (10°N—21°N) 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 resembling the
Australian taxon, but the flattened branches of R. usnea do not occur in R. filicaulis. There are
morphological similarities between the Canary Island species R. chondrina and the Australian
RAMALINA IN AUSTRALIA 161
taxon; both are subtropical species which occur in the mountains. Krog & Msthagen (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-like, often minute; up to 1-0 cm high; branching dense and intricate, distally producing
branchlets; branch width 0-3—1-0 mm, branches flat to subterete, variably inflated, some
branches with slit-like perforations; surface shiny to matt, smooth; holdfast diffuse; soralia form
from disintegration of the lower cortex occurring laminally and at the apices (Plate 1, fig. 5).
Apothecia uncommon, but numerous on some thalli, lateral or terminal, disc 0-5—3-0 mm
diam., concave, plane or convex; margin entire or incised at maturity; spores ellipsoid, straight
or rarely curved, 8-13 x 4-5 um.
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 Australia this
species is saxicolous only, although in East Africa it is both saxicolous and corticolous (Krog &
Swinscow, 1974).
Its occurrence on several types of rock (viz. granite, sandstone, conglomerate, trachyte, and
dolerite) shows it is not substrate specific. In some localities R. fimbriata occupies sheltered
overhangs and ledges, habitats which would have little moisture available for the thalli, apart
162 G. NELL STEVENS
120 130 140 160
10N
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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, JAE (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 & J C (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 Mill. 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 Mill. 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. Wilsons.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. Péytiak. 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—)1—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) um.
Chemistry. Sekikaic acid (+ trace homosekikaic acid or trace divaricatic acid), or usnic acid
only, or divaricatic acid and usnic acid.
Remarks. The type material of R. glaucescens comprises a branch covered with tiny tufted
specimens and a card with four specimens attached to it (which are here designated a, b, c, d).
Two growth forms are represented on the card; a and d have bifurcate branches with small
apothecia lying at the axil of the fork, and b and c have broader branches and bear terminal or
subterminal apothecia, the latter with small subtending spurs. All of the thalli tested contain
sekikaic acid. The specimens are exceptionally small and are immature.
The considerable polymorphism encountered in R. glaucescens presented much difficulty in
interpretation, as there was no correlation with the chemical variation; a situation rather similar
to that encountered by Kristinsson (1969) in dealing with Cetraria islandica in Iceland. Five
different species names already existed for entities now included in the complex: R. glaucescens
Krempelh., R. leiodea var. fastigiatula Mill. Arg., R. brevis F. Wilson, R. lacerata Mill. 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
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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
1230 130 140 160
1 1 1 n
Fig. 13 Distribution of R. glaucescens: @ sekikaic acid race; O acid-deficient race; * divaricatic acid race.
or a particular bark substrate has an effect on this phenotype’s distribution. Sekikaic acid was
always present in such thalli.
One ecotype of R. glaucescens grows along the southern Victorian coastline; it has a short
broad thallus with large terminal or laminal apothecia and is acid-deficient. A few specimens
resembling this morphology have been collected from inland in eastern Australia, but that
material contains sekikaic acid and is not so coarsely textured as the coastal thalli. This ecotype
resembles the type of R. brevis (Plate 8, fig. 15)..
Another acid-deficient ecotype occurs inland at low altitudes in Victoria and South Australia,
but the thallus is generally narrower than that of the coastal ecotype and the apothecia are not as
large; however, intergradation between the two is evident at some locations (Plate 8, fig. 8). The
presence of small branchlets on the main branches resembles the eastern highlands ecotype. The
type of R. calicaris var. australica belongs to this ecotype. In some habitats the thallus becomes
quite coarse in texture with chondroid strands showing at the surface; such characteristics are
apparent in the material from the arid zone in Western Australia, including the type collection of
R. lacerata.
The largest morphotype in the R. glaucescens complex is found in western Victoria and
eastern South Australia, with a few specimens collected in western New South Wales. The latter
specimens contain sekikaic acid but all other material proved to be acid-deficient. Most of the
ecotypes of R. glaucescens have been found to intergrade with this large morphotype (Plate 8,
fig. 1). It is normally sterile although pycnidia have been detected in some of these specimens.
Four of the type specimens here included in R. glaucescens were initially collected within 100
RAMALINA IN AUSTRALIA 167
km radius of one another in Victoria, which illustrates the ecophenotypic response to different
habitats. Table 9 summarises the morphological variation that is found in the different ecotypes.
This taxon is entirely corticolous, the most common phorophytes being Acacia spp. and
Bursaria spinosa. It occurs as dense colonies along the branches of isolated trees in pastureland
or in open-forest communities. In some localities the entire population may be small and
immature (as found in the type material of both R. glaucescens and R. leiodea var. fastigiatula),
whereas in other areas only large mature specimens occur.
World distribution. Australia and New Zealand.
New South Wales: Ben Lomond, 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 Leptospermumsp..,
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 Pinus 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,
GNS (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, RF (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, GB & JC (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);
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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, RWR (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, J. Oliver (G, M). Tasmania: Trowutta, 41°02’S, 145°05’E, on Malus sp., 1981,
GK 384/81; Beaconfield, 41°12’S, 146°45’E, on Acacia dealbata, 1980, GK 214/80; Prossers Forest Rd,
41°15'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, GK 31/80.
8. Ramalina litorea 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 wm.
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, rhyolitic, 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 I, 23°10’S, 150°58’E, substrate unknown, 1976, U. Allen (H032240); Double Head, 23°08’S,
150°47’E, on slates (15 melev.), 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
120 130 140 150
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9. Ramalina peruviana Ach. Plate 9, figs 1-2.
Lichenogr. Univ.: 599 (1810). Type: South America, Peru, Lagastas.n. (H-ACH! — holotype; BM!, UPS!
— isotypes; sekikaic acid aggregate, and usnic acid).
Ramaliria farinacea var. dendroides Mill. 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 Mill. 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).
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RAMALINA IN AUSTRALIA
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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 wm.
Chemistry. Homosekikaic acid (M), sekikaic acid (M), ramalinolic acid (m/t), 4’-0-
demethylsekikaic acid (m/t), 4’-O-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 (Landr6n, 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.
Jgrgensen (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 litt.).
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 173
120 130 140 150
30
40
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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 celaoxyloides, 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 I. 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 Acacia sp., 1976, R. Moodie (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 I. 32°56’S, 151°46’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).
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, Tapper 933 (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) um.
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
Australia.
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
1 T
120 130 140 150
1 n i i
Fig. 16 Distribution of R. reducta.
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 from 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 thalli 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 Banksia sp., 1982, GNS (BRIU3632NS); on Acacia 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 Mill. 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, delicate, 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, linear
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 the material from latitudes 21°S—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 77
¥ 7 ui m ip
120 130 140 150
a 104
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e
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9
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Fig. 17 Distribution of R. tenella.
var. confirmata, R. pacifica, R. exiguella, R. nervulosa var. nervulosa, and R. nervulosa var.
luciae.
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, GNS (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 « 4-4-5 wm.
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 of R. 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 R. 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°0S’E, on Hakea sp., 1976 JAE (JAE 2261); Mt Barker, 35°06’S,
138°51'E, on dead twig, 1981, RWR (BRIU2427RR). Tasmania: Trowutta, 41°02’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, 147°44’E, on Acacia dealbata, 1981, GK 60/81; Cressy, 41°43’S, 147°02'E,
on dead Acacia, 1964, JC (H032561); Central Plateau, 41°45’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, GB & JC (H032557); Dee, 6km 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; Ellendale, 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 (H032330); 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 [‘whinrayi? ] 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 wm.
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 R. 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,
180 G. NELL STEVENS
a Bia
120 130 140 160
104
20~
30+
40%
L | \ t
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 B 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
Plate10 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
TTTITITIFIDT eee
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 (1—)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 wm.
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, 19835). Broad thalli which
belong to R. celastri subsp. celastri resemble the morphology of the holotype of R. yemensis
(Ach.) Nyl. (R. fraxinea B 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 of R. yemensis.
Weber & Wetmore (1963, 1972) listed the names R. ecklonii Sprengel, R. ecklonii var.
membranacea (Laurer) Mill. Arg., R. ecklonii var. tenuissima Meyen & Flotow, R. linearis Sw..,
and R. yemensis (Ach.) Nyl. as occurring in Australia, 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 Fée 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 of R. celastri from Mt Kenya resembled R. cumanensis, and Landroén (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 and 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 A. 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 183
120 130 140 160
10+
30~
nN l
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 Acacia sp., 1978, GNS (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); Wollomombi 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, GK 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, Gunns.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 wm.
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
Australia, and Western Australia 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 litt.) detected the occurrence of this broad taxon in collections of R. celastri from East
Africa. Examination of the BM material from Tanzania, Moshi district (1440 m alt.) and
Uganda (1950-2100 m alt.) showed that thalli 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 of R. 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: Marlo, 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°5S0’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, J. H. Willis (MEL 10025); Long I, 34°03’S, 121°58’E, on
Melaleuca sp., 1950, J. 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 Bursaria spinosa, 1965, J. 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 & JC (H032314).
15. Ramalina inflata (J. D. Hook. & Taylor) J. D. Hook. & Taylor
in J. D. Hook., Flora Antarctica 1: 194 (1845).
Cetraria inflata J.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. inflata Plate 10, figs 5—6; Plate 14, fig. 5.
Ramalina tasmanica Ny). in Bull. Soc. Linn. Normandie II, 4: 162 (1870) type: Australia, Tasmania, J. 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 1—-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) wm.
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
lichenologists (Howe, 1913-14; Landr6én, 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 of R. 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 &
@sthagen (1980).
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
T
120 130 140 160
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Fig. 21 Distribution of R. inflata subsp: inflata O, subsp. perpusilla @, and subsp. australis ®.
RAMALINA IN AUSTRALIA 187
120 130 140 1bo
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Fig. 22 Distribution of R. inflata complex: @ divaricatic acid race; O sekikaic acid race.
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 (Wollomombi 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);
Wollomombi 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 Casuarina sp. n., 1978, JAE (JAE 4543); Spring Ck
188 G. NELL STEVENS
Gap, 37°30’S, 148°20’E, on Acacia sp., 1978, JAE (JAE5156); Marlo, 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); Healesville, 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, 41°09’S, 145°40’E, on Malus sp., 1980, GNS
(BRIU3239NS); Mt Barrow, 41°23’S, 147°25’E, on Acacia sp., 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 Mill. 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,
1—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 um.
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 would 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, €
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
many of the thalli.
The morphology of the East African taxon R. consanguinea Mill 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) wm, 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, GNS (BRIU4114NS); Burdekin R, 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, I. 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, 151°45’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 Citriobatus pauciflora, 1979, GNS (BRIU3654NS); Stradbroke I, 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, GNS (BRIU3010NS); Mt Cordeaux, 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,°GNS (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. Beallingall
(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. australis N. 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) wm. 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 laminal; 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) wm.
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 & @sthagen (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 Australia 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 thalli
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 small
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 of R. inflata subsp. australis.
G. NELL STEVENS
192
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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, and 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: Marlo, 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, RF (MEL1026193); Drik Drik,
38°00’S, 141°18’E, 1952, A. 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, J. Z. Weber (AD97647391); Naracoorte, 25 km SE of, 36°58’S, 140°45’E, 1969, J.
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, J. H. Willis (MEL1026176); Yalata Road
House, 5 km W of, 31°39’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 67/366; 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, 41°13’S, 146°45’E, on Acacia dealbata, 1980, GK 216/80;
Buckland, Pitts Hill, 42°36’S, 147°43’E, on Bursaris spinosa, 1980, GK 450/80; Hobart, Gunners Quoin,
42°54’S, 147°19’E, on dolerite rock, 1980, GK 600/80.
16. Ramalina fissa (Mill. Arg.) Vainio Plate 10, fig. 8.
in Mem. Herb. Boissier 5: 2 (1900).
Ramalina inflata var. fissa Mill. 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: Australia,
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; soralia absent.
Apothecia numerous, terminal, not spurred; sometimes laminal; disc 2—5(—8) mm diam.,
initially concave becoming flat; margin entire, inrolled; spores ellipsoid, mainly curved, some
straight, (10—)12—16 x 4—5(-6) um.
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
of R. fissa.
Wilson identified large coarse morphotypes of this taxon as R. fraxinea f. platyna Ny].
Material held at G under the latter name consists of two thalli 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. inflata) 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. inflata 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 thalli 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 of R. 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 Marlo, Victoria.
It does not show any substrate preference.
World distribution. Australia.
Victoria: Linga, 13 km N of, Pink Lakes, 35°06’S, 141°40’E, on dead wood, 1970, GB & JC (H032663);
Casterton, 51-5 km N of, 37°20’S, 141°20’E, on Acacia sp., 1973, RF (MEL1036009); Creswick, 37°26'S,
143°54’E, on Pinus sp., 1963, J. H. Willis (MEL9335); Lake Burrumbeet, 37°28’S, 143°40'E, on Pinus
sp., 1945, P. Bibby (MEL10018); Marlo, 37°48’S, 148°42’E, on fence post, 1978, JAE (JAES195);
Troidea Hill, 37°57’S, 141°19’E, on Banksia sp., 1965, RF (MEL1026193); Portsea, 38°20’S, 144°42’E,
on shrub, 1964, J. Williams (MEL1516513); Portland, 38°21’S, 141°36’E, on Malus 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, GNS (BRIU3198NS).
Bass Strait: Hogans I, 39°13’S, 146°59’E, on Leucopogon sp., 1973, JW (MEL10130013); Little Dog i,
RAMALINA IN AUSTRALIA 195
130 130 140 150
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Fig. 23 Distribution of R. fissa.
40°15'S, 148°17’E, on twigs, 1977, JW (MEL1026160); Flinders I, 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
(HO32296); 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). South
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, RF (MEL1020693).
17. Ramalina nervulosa (Mill. Arg.) des Abb.
in Bull. Inst. fr. Afr. noire 14: 25 (1952).
Ramalina farinacea var. nervulosa Mill. 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—)1-0—1-5(—2-0) mm,
196 G. NELL STEVENS
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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 ellipsoid. Apothecia not seen.
Chemistry. Divaricatic acid and stenosporic acid and usnic acid.
Remarks. Miller [Arg.] (1883) applied 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 (Mill. Arg.) des Abb. (Stevens, 1983a).
When the five specimens held at G in the Miller [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 Miller (1883) only the Australian material from Daintree River Queensland
contains divaricatic acid and stenosporic acid. This specimen was designated as lectotype by
Filson in 1971.
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); Barrons 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—1-0(—
< 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
5 104
, SR ; _, 800km WS,
as
Ee Nas ak a a a
\&
_ 30
oH
L LS 404
1 Ll 1 1
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 delimited; 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 tlc] 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 R. nervulosa var. dumeticola in Australia at
subtropical latitudes appears to be an anomaly (Fig. 25). Previous research (Stevens, 1983a) 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
t tT Li Li
420 130 140 160
. S? 1O-
| a 1 1 ie i: _890km
CQ
e@
@
é 20
WN *
0
ro
*
f 304
oe
(oe)
$
ie 40~
1 4 ue L
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:
Chaelundi Mt, 37 km N of Ebor, 30°04’S, 152°21'E, Acacia orrites, 1978, D. Verdon (CBG7809394); Cattle
Ck State Forest, 12 km NNE of Dorrigo, 30°15’S, 152°03’E, on Banksia integrifolia, 1978, D. Verdon
(CBG8203532).
var. luciae (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—1-0(—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) um.
Chemistry. Sekikaic acid (M), 4’-0-methylnorsekikaic acid (m), 4’-0-demethylsekikaic acid (m
or t), homosekikaic acid (t), ramalinolic acid (t), 4’-O-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 R.
farinacea var. nervulosa Mill. Arg., was collected from Tahiti last century by Jardin. This taxon
is acommon 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, 1983a).
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 of R. 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 Australia 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,
1983a).
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 I, 18°17’S, 146°13’E, on Ceriops tagal, 1979, GNS (BRIU3137NS); Eimeo,
21°09'S, 149°09’E, on Rhizophora stylosa, 1976, GNS (BRIU3542NS); St Lawrence, 22°21’S, 149°32’E, on
Ceriops tagal, 1980, GNS (BRIU3600NS); 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, GNS (BRIU280NS); Toogoom, 25°15’S, 152°40’E, 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 pacifica Asah. Plate 11, fig. 1; Plate 14, fig. 8.
in J. 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—)1—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 revolute.
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 um.
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 thalli 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 insular'um 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
Australia, 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 thalli are generally
broader than the thalli from the higher rainfall habitats. Only one inland collection of R. 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
cs rE T
120 130 140 160
i 104
o 800km ST
oe ry . ) 1 i
e
e
a e
20
oF
4,
ys
30
oe
os
ie ie 404
L 1 i 1
Fig. 26 Distribution of R. pacifica.
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 established on the cliff face.
World distribution. Burma/Bangladesh, Japan, Indonesia (Java), Marianas Is, South Africa,
Australia, 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, GNS (BRIU3284NS); Shute Harbour, 20°16’S, 148°47’E, on Ceriops tagal, 1979,
GNS (BRIU3283NS); Brampton I, 20°49’S, 149°18’E, J. M. Gilbert (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, 151°16’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, 151°40’E, on Ceriops tagal,
1975, GNS (BRIU1153NS); Round Hill Hd., 24°10’S, 151°54’E, on Rhizophora stylosa, 1975, RWR
(BRIU915RR); Fraser 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, GNS (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 subfraxinea Ny.
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 1 es7; Plate 42; fies3,. 7; Plate 15, fies 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) um.
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 (Gretan 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°51'S 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,
u
120 130 140 160
L 1 1 i
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, J. 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. Ballment (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, 151°40’E, on Rhizophora stylosa, 1975, GNS (BRIU1278NS); Turkey, 24°06’S, 151°37’E, on
Rhizophora stylosa, 1975, GNS (BRIU1150NS).
var. confirmata (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 Il, 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 um.
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 litt.) 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 I., north of Cape York) to latitude 31°S
(Fig. 28).
Although mostly found along the shoreline, R. subfraxinea var. confirmata has been collected
from the coastal ranges at Montville, Kenilworth, 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
' ak
120 130 140 e 150
9
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i \
{ |
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 Indian.
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 I, 14°42’S, 145°11’E, on
beach shrub, 1979, J. 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
Lumnitzera sp., 1979, GNS (BRIU3465NS); Townsville, 19°15’S, 146°50’E, on Rhizophora stylosa, 1975,
B. Ballment (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, 151°16’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); Bribie I, 27°02’S, 153°08’E, on Lumnitzera racemosa, 1978, GNS (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); Coochiemudlo I, 27°32'S, 153°17’E, on Callitris columel-
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, RWR (BRIU3883*NS); Yamba, 29°27’S, 153°20’E, on Avicennia marina, 1977, GNS
(BRIU3177NS); Port Macquarie, 31°27'S, 152°54’E, on Ficus sp., 1975, JAE (JAE 1089); Hunter R., Ash
1,32°56'S,-151°46'E, 1845;.J2P. Verreaux (PC):
var. leiodea (Nyl.) N. Stevens, comb. nov. Plate 11, figi6; Plate 12, fio.;8; Plate 15; fie 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, ‘Donné par le Musée Colonial’,
1861, Deplanche s.n. (PC! — lectotype; boninic acid aggregate).
Ramalina boninensis Asah. in J. 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 J. Jap. Bot. 15: 211 (1939). Type: Bonin Islands,
Ogasawara I., Hahajima, 1936, Kusakas.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 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 to plane; margin entire or
incised at maturity; spores ellipsoid, straight or slightly curved; 10—12(—16) x 4-5 um.
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 wm, whereas R. maritima spores lie in the
range 18-25 x 5-6 um.
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, Réunion I., Australia, 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 (BRIU3225N§); 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
I, 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,
GNS (BRIU4311NS).
var. norstictica N. 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 um.
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; soralia 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) Xx 3-5—5-5 wm.
Chemistry. Norstictic acid, + connorstictic acid, and usnic acid.
Remarks. This taxon is an Australian endemic; examination of esorediate maritime taxa from
overseas herbaria failed to find any norstictic acid taxon which resembled it. In Australia 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 vat. 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
Australian coastline (Figs 29, 31).
Distribution and habitat. Ramalina subfraxinea var. norstictica occurs along the northern and
RAMALINA IN AUSTRALIA 209
, A T T
120 130 140 150
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Fig. 29 Distribution of R. subfraxinea var. norstictica.
eastern coastlines of Australia (Fig. 29). Its absence from an area between latitudes 17°-11°S 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
thalli 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, J. 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.
Ballment (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,
RWR (BRIU1039RR); Gladstone, 23°50’S, 151°16’E, on Rhizophora stylosa, 1980, GNS (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, 11-18(-—22) x 3-8-S(-6) wm. 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 along curved spur up to 10 mm, disc (1—)1-5—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) wm.
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 canaliculate
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 R. subfraxinea var. leiodea. Fig. 9 R. subfraxinea var. norstictica. Fig. 10 R. subfraxinea var.
confirmata.
RAMALINA IN AUSTRALIA
212 G. NELL STEVENS
120 130 140 150
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204
30
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Fig. 30 Distribution of R. tropica.
Distribution and habitat. The distribution of R. tropica is restricted to an area on the eastern
coastline 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 thalli.
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
Tropic
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 Molloy, 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 I, 18°20’S, 146°15’E, on Ceriops tagal, 1979, GNS (BRIU3160NS).
214 G. NELL STEVENS
BERS. MUS. Panis,
| PH ays re
ele Sp )5
‘oH OBIE MULLER
4V7O9/
Dn (Brwroar.e front APA re b> Ui
ty fin Fi MA lor. Nrrevhan FFF
Plate 13. Photographs of Australian 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 var. fastigiatula (G). Fig. 7 R. calicaris var. australica (H). Fig. 8 R. brevis (NSW).
RAMALINA IN AUSTRALIA 215
Ps Pe
AD ci a Ce
(i PIER MULLER ANB. 1896 Karn atin Du / Bey aeroree.
thallus, Kem: KC -PD
medulia: Kee Commi Cae PD potent! 4%. Kimurhk Ses vt.
Plate 14 Photographs of Australian Ramalina types. Scale in mm. Fig. 1 R. farinacea var. squarrosa (G).
Fig. 2 R. farinacea var. dendroides (G). Fig. 3 R. 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 R. pacifica (TNS).
216
G. NELL STEVENS
GS2 Vea nahaa Flee fo. i dow
Shenk. — i gy, fiatgann ptr tads
fh. fo ther PRM Woden
— pe
os NE at
pass Kye th Les ach.
a ers
2 OkBrs Sh ed save g 7593,
RAISUAAING GUVANVE "WAIN VENSH MOTH TE
eg oo
Pia te
- Liotta
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 Australia in Wetmore (1963), and Weber & Wetmore
(1972) is indicated below.
. anceps = R. australiensis Nyl. or R. filicaulis N. Stevens
. angulosa = R. australiensis Nyl.
. calicaris = R. glaucescens Krempelh.
. calicaris var. canaliculata = R. glaucescens Krempelh.
. complanata = R. subfraxinea Ny].
confirmata = R. subfraxinea var. confirmata (Nyl.) N. Stevens.
dendriscoides = R. tenella Mill. Arg. or R. peruviana Ach.
dendriscoides var. minor = R. tenella Mill. Arg. or R. peruviana Ach.
. dilacerata var. alba = R. inflata subsp. perpusilla (Stirton) N. Stevens
ecklonii = R. celastri (Sprengel) Krog & Swinscow
ecklonii var. membranacea = R. celastri (Sprengel) Krog & Swinscow
ecklonii var. ovalis = R. celastri subsp. ovalis (J. D. Hook & Taylor) N. Stevens
ecklonii var. tenuissima = R. celastri (Sprengel) Krog & Swinscow
. fastigiata = R. glaucescens Krempelh.
fraxinea = R. glaucescens Krempelh.
fraxinea var. ampliata = R. glaucescens Krempelh.
. fraxinea f. platyna = R. fissa (Mill. Arg.) Vainio
. fraxinea var. taeniata = R. glaucescens Krempelh.
furcellata = R. australiensis Ny).
geniculata = R. inflataJ. D. Hook. & Taylor
geniculata var. olivacea = R. inflata subsp. perpusilla (Stirton) N. Stevens
gracilenta = R. exiguella Stirton
gracilis = R. exiguella Stirton or R. filicaulis N. Stevens
homalea = unknown?
inflata var. gracilis = R. inflata (J. D. Hook. & Taylor) J. D. Hook. & Taylor
intermedia = R. inflata subsp. australis N. Stevens
javanica = R. peruviana Ach.
linearis = R. celastri (Sprengel) Krog & Swinscow
. pollinaria = R. unilateralis F. Wilson
polymorpha = unknown
polymorpha f. emplecta = unknown
pusilla = R. inflata subsp. australis N. Stevens
scopulorum = R. subfraxinea Ny]. var. ?
scopulorum var. cuspidata = R. subfraxinea Nyl. var. ?
scopulorum var. subfarinacea = R. pacifica Asah.
. tayloriana = R. nervulosa var. luciae (Molho et al.) N. Stevens
usnea = R. australiensis Nyl.
. usneoides = R. australiensis Nyl.
. yemensis = R. celastri (Sprengel) Krog & Swinscow
. yemensis var. ovalis = R. celastri subsp. ovalis (J. D. Hook. & Taylor) N. Stevens
DD WWD DDD DDD D DD DAD DDR DDD RD DDR WD DR DDD RAZ
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 Ramalinae), 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 Ramalinae), 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.
References
Abbayes, H. des 1952. Lichens récoltés en Guinée Francaise et en Céte d'Ivoire V-VI. Bull. Inst. fr. Afr.
noire B, 14: 19-27.
Acharius, E. 1810. Lichenographia universalis. Gottingen.
— 1814. Synopsis methodica lichenum. Lund.
Agardh, C. A. 1821. Aphorismi botanici. Lund.
Asahina, Y. 1938. Lichenologische Notizen. X. J. Jap. Bot. 14: 251-255.
— 1939. Ramalina-Arten aus Japan (II). J. Jap. Bot. 15: 205-223.
Awasthi, D. D. 1965. Catalogue of the lichens from India, Nepal, Pakistan and Ceylon. Beih. Nova
Hedwigia 17: 1-137.
Bailey, F. M. 1886. A synopsis of the Queensland flora containing both the phaenogamous and cryptog-
amous plants. First supplement: 1—99. Brisbane.
Beltman, H. A. 1978. Vegetative Strukturen der Parmeliaceae und ihre Entwicklung. [Bibltheca lich. 11].
Vaduz.
Bowler, P. A. 1977. Ramalina thrausta in North America. Bryologist 80: 529-532.
— 1981. Cortical diversity in the Ramalinaceae. Can. J. Bot. 59: 427-452.
& Rundel, P. W. 1972a. Ramalina bajacalifornica. A new species of lichen from Baja California.
Bryologist 75: 365-366.
— 1972b. A new species of Ramalina from Sinaloa, Mexico. Bryologist 75: 574-576.
—— — 1973. Two new lichens (Ramalina) from Baja California, Mexico. Bryologist 76: 211-213.
—— — 1974. The Ramalina intermedia complex in North America. Bryologist 77: 617-623.
—— —— 1977. Synopsis of a new lichen genus Fistulariella Bowler & Rundel (Ramalinaceae). Mycotaxon
6: 195-202.
1978. The Ramalina farinacea complex in North America: chemical, ecological and morpho-
logical variation. Bryologist 81: 386—403.
Brandt, T. 1906. Beitrage zur anatomischen Kenntnis der Flechtengattung Ramalina. Hedwigia 45:
124-158.
Brodo, I. M. 1978. Changing concepts regarding chemical diversity in lichens. Lichenologist 10: 1-11.
—— & Hawksworth, D. L. 1977. Alectoria and allied genera in North America. Op. bot. Soc. bot. Lund 42:
1-164.
Carey, S. W. 1976. The expanding Earth. Amsterdam.
Chester, D. O. & Elix, J. A. 1978. The identification of four new metadepsides in the lichen Ramalina
asahinae. Aust. J. Chem. 31: 2745-2749.
Choisy, M. 1954. Classification des lichens fruiticuleux. Bull. Mens. Soc. linn. Lyon 23: 229-240.
Croizat, L. 1952. Manual of phytogeography. Den Haag.
, Nelson, G. & Rosen, D. E. 1974. Centres of origin and related concepts. Syst. Zool. 23: 265-287.
Crombie, J. M. 1880. Enumeration of Australian lichens in Herb. Robert Brown (Brit. Mus.), with
descriptions of new species. J. Linn. Soc. Bot. 17: 390-401.
Culberson, C. F. 1972. Improved conditions and new data for the identification of lichen products by a
standardized thin-layer chromatographic method. J. Chromat. 72: 113-125.
—— & Culberson, W. L. 1977. Chemosyndromic variation in lichens. Syst. Bot. 1: 325-339.
—— & Hale, M. E. 1973. Chemical and morphological evolution in Parmelia sect. Hypotrachyna. Product
of ancient hybridization? Brittonia 25: 162-173.
Culberson, W. L. 1967. Analysis of chemical and morphological variation in the Ramalina siliquosa species
complex. Brittonia 19: 333-352. 5
—— 1969a. The behaviour of the species of the Ramalina siliquosa group in Portugal. Ost. bot. Z. 116:
85-94.
—— 1969b. The use of chemistry in the systematics of the lichens. Taxon 18: 152-166.
—— & Culberson, C. F. 1967. Habitat selection by chemically differentiated races of lichens. Science,
N. Y. 158: 1195-1197.
—— —— 1968. The lichen genera Cetrelia and Platismatia (Parmeliaceae). Contr. U.S. natn. Herb. 34:
449-558.
RAMALINA IN AUSTRALIA 219
—— —— 1970. A phylogenetic view of chemical evolution in the lichens. Bryologist 73: 1-31.
—_—-—— & Johnson, A. 1977. Correlation between secondary-product chemistry and ecogeography in the
Ramalina siliquosa group (lichens). Plant Syst. Evol. 127: 191-200.
Ducker, S. C. 1979. History of Australian phycology: the significance of early French exploration.
Brunonia 2: 19-42.
Duncan, U. K. 1970. Introduction to British Lichens. Arbroath.
Du Rietz, G. E. 1924. Die Soredien und Isidien der Flechten. Svensk. bot. Tidskr. 18: 371-396.
— 1926. Morfologi och systematik hos slaktet Ramalina sarskilt dess skandinaviska arter. Svensk. bot.
Tidskr. 20: 295-299.
Elix, J. A. 1982. Peculiarities of the Australasian lichen flora: accessory metabolites, chemical and hybrid
strains. J. Hattori bot. Lab. 52: 407-415.
Eschweiler, F. G. 1824. Systema lichenum genera exhibens rite distincta pluribus novis adaucta. Niirnberg.
Filson, R. B. 1976. Australian lichenology: a brief history. Muelleria 3 (3): 183-190.
—— & Rogers, R. W. 1979. Lichens of South Australia. Adelaide.
Follmann, G. 1967. Die Flechtenflora der nordchilenischen Nebeloase Cerro Moreno. Nova Hedwigia 14:
215-281.
— & Huneck, S. 1969. Mitteilungen iiber Flechteninhaltsstoffe LXI. Zur Chemotaxonomie der
Flechtenfamilie Ramalinaceae. Willdenowia 5 (2): 181-216.
Galloway, D. J. 1979. Biogeographical elements in the New Zealand lichen flora. Jn D. Bramwell (Ed.),
Plants and islands: 201-224. London.
— 1985. Flora of New Zealand. Lichens. Wellington.
Hale, M. E. 1973. Fine structure of the cortex in the lichen family Parmeliaceae viewed with the scanning
electron microscope. Smithson. Contr. Bot. 10: 1-92.
— 1975. A monograph of the lichen genus Relicina (Parmeliaceae). Smithson. Contr. Bot. 26:
1-32.
— 1976a. A monograph of the lichen genus Pseudoparmelia Lynge (Parmeliaceae). Smithson. Contr.
Bot. 31: 1-62.
— 1976b. A monograph of the lichen genus Bulbothrix Hale (Parmeliaceae). Smithson. Contr. Bot. 32:
1-29.
— 1976c. A monograph of the lichen genus Parmelina Hale (Parmeliaceae). Smithson. Contr. Bot. 33:
1-60.
— 1978. A new species of Ramalina from North America (Lichenes: Ramalinaceae). Bryologist 81:
599-601.
1981. Pseudocyphellae and pored epicortex in the Parmeliaceae: their delimitation and evolutionary
significance. Lichenologist 13: 1-10.
& Culberson, W. L. 1970. A fourth checklist of the lichens of the continental United States and
Canada. Bryologist 73: 499-543.
Hawksworth, D. L. 1976. Lichen chemotaxonomy. Jn D. H. Brown, D. L. Hawksworth, & R. H. Bailey
(Eds), Lichenology: progress and problems: 139-148. London.
— 1977. A bibliographic guide to the lichen floras of the world. Jn M. R. D. Seaward (Ed.) Lichen
ecology: 437-502. London.
Henssen, A. & Jahns, H. M. 1973 [‘1974’]. Lichenes. Stuttgart.
Holmgren, P. K., Keuken, W. & Schofield, E. K. (Eds) 1981. Index herbariorum. 7th ed. [Regnum veg.
106].
Hooker, J. D. & Taylor, T. 1844. Lichenes antarctici; being characters and brief descriptions of the new
lichens discovered in the southern circumpolar regions, Van Diemen’s Land and New Zealand, during
the voyage of H.M. discovery ships Erebus and Terror. J. Bot. Lond. 3: 634-658.
Howe, R. H. 1913-14. North American species of the genus Ramalina I-VIII. Bryologist 16: 65—74, 81—89;
17: 1-7, 17-27, 33-40, 49-52, 65-69, 81-87.
Hue, A. M. 1890. Lichenes exoticos. Nouv. Archs Mus. Hist. nat. Paris U1, 2: 209-322.
1901. Lichenes extra-Europaei, a pluribus collectoribus ad museum Parisiense missi. Nouv. Archs
Mus. Hist. nat. Paris, V1, 3: 80-122.
Imshaug, H. A. 1972. Typification of Ramalina usnea (L.) R. H. Howe. Lichenologist 5: 317-318.
— & Brodo, I. M. 1966. Biosystematic studies on Lecanora pallida, and some related lichens in the
Americas. Nova Hedwigia 12: 1-59.
Jahns, H. M. 1974 [‘1973’]. Anatomy, morphology and development. Jn V. Ahmadjian & M. E. Hale
(Eds), The lichens: 3-58. New York.
Jergensen, P. M. 1974. Dirinaria applanata (Fée) Awasthi on the Agores and a comment on the
taxonomical value of sekikaic acid in the genus. Port. Acta biol. B, 12, 5-9.
220 G. NELL STEVENS
1977. Foliose and fruticose lichens from Tristan da Cunha. Norske vidensk.-Akad. I. Mat.-
Naturvitensk. Kl. Skr. 11, 36: 1-40.
1979. Phytogeographical relationship of the lichen flora of Tristan da Cunha (excluding Gough
Island). Can. J. Bot. 57: 2279-2282.
— 1983. Distribution patterns of lichens in the Pacific region. Aust. J. Bot., supp. ser. 10: 43-66.
Keuck, G. 1979. Die systematische stellung der Ramalinaceae. Ber. dt. bot. Ges. 92: 507-518.
Kristinsson, H. 1969. Chemical and morphological variation in the Cetraria islandica complex in Iceland.
Bryologist 72: 344-357.
Krog, H. & James, P. W. 1977. The genus Ramalina in Fennoscandia and the British Isles. Norw. J. Bot.
24: 15-43.
— & Osthagen, H. 1980. The genus Ramalina in the Canary Islands. Norw. J. Bot. 27: 255-296.
— & Swinscow, T. D. V. 1974. Ramalina species with a hollow thallus (Fistularia) in East Africa. Norw.
J. Bot. 21: 111-124.
— — 1975. Some Ramalina species with punctiform pseudocyphellae in East Africa. Norw. J. Bot. 22:
269-276.
—— — 1976. The genus Ramalina in East Africa. Norw. J. Bot. 23: 153-175.
Landron, C. I. 1972. The lichen genus Ramalina Ach. in the West Indies with notes on its role in the
vegetation of Puerto Rico. Ph.D. thesis (unpublished), Michigan State University.
Magnusson, A. H. 1956. A catalogue of the Hawaiian lichens. Ark. Bot. II, 3: 223-402.
—— & Zahlbruckner, A. 1945. Hawaiian Lichens III. The families Usneaceae and Physciaceae. Ark. Bot.
32A (2): 1-89.
Malme, G. O. 1934. Die Ramalinen der ersten Regnellschen Expedition. Ark. Bot. 26A (12): 1-9.
Massalongo, A. B. 1854. Neagenea lichenum. Verona.
Moberg, R. 1977. The lichen genus Physcia and allied genera in Fennoscandia. Symb. Bot. Upsal. 22 (1):
1-108.
Molho, D., Bodo, B., Culberson, W. L. & Culberson, C. F. 1981. A chemically distinctive new Ramalina
from Fiji. Bryologist 84: 396-398.
Montagne, J. F. C. 1852. Lichens. Jn C. Gay, Historia fiscia y politica de Chile 8: 53-228.
Moore, B. J. 1968. The macrolichen flora of Florida. Bryologist 71: 161-266.
Miller, J. 1883. Lichenologische Beitrage XVIII. Flora, jena 66: 17-25.
Nix, H. A. 1981. The environment of Terra Australis. Jn A. Keast (Ed.), Ecological biogeography of
Australia: 105-133. Den Haag.
— 1982. Environmental determinants of biogeography and evolution in terra australis. Jn W. R. Barker
& P. J. M. Greenslade (Eds), Evolution of the flora and fauna of arid Australia: 47-64. Frewville.
Nylander, W. 1857. Enumération générale des lichens, avec l’indication sommaire de leur distribution
géographique. Mém. Soc. Sci. nat. Cherbourg 5: 85-146.
— 1859. Prodromus expositionis lichenum Novae Caledoniae. Annis Sci. nat. (Bot.) IV, 14: 280-283.
—— 1861. Expositio lichenum Novae Caledoniae. Annals Sci. nat. (Bot.) IV, 15: 37-54.
—— 1870. Recognitio monographica Ramalinarum. Bull. Soc. Linn. Normandie I1, 4: 101-181.
—— 1888. Lichenes Novae Zelandiae. Paris.
Osorio, H. S. 1970a. Contribution to the lichen flora of Argentina IV. New or additional records.
Bryologist 73: 392-394.
1970b. Lichenes from Cantera, South Paraguay. Comun. bot. Mus. Hist. nat. Montev. 4 (50): 1-3.
— 1972. Contribution to the lichen flora of Uruguay VII. A preliminary catalogue. Commun. bot. Mus.
Hist. nat. Montev. 4 (56); 1-46.
1978. Contribution to the lichen flora of Brazil IV. Lichens from southern Rio Grande do Sul.
Bryologist 8: 452-454.
, Aguiar, L. W. & Zanette, V. C. 1980. Contribution to the lichen flora of Brazil VII. Lichens
from Montenegro and Triunfo, Rio Grande do Sul state. Comun. bot. Mus. Hist. nat. Montev. 4 (62):
1-8.
Peveling, E. 1970. Die Darstellung der Oberflachenstrukturen von Flechten mit dem Raster-
Elektronenmikroskop. Ber. dt. bot. Ges. II, 4: 89-101.
—— 1974 [‘1973’]. Fine structure. Jn V. Ahmadjian & M. E. Hale (Eds), The lichens: 147-182. New York.
Poelt, J. 1969. Ramalina. Bestimmungsschliissel Europdischer Flechten: 550-562. Lehre.
— 1970. Das Konzept der Artenpaare bei den Flechten. Vortr. GesGeb. Bot. II, 4: 187—198.
—— 1972. Die taxonomische Bedhandlung von Arten-paaren bei den Flechten. Bot. Notiser 125: 77-81.
— 1974 [‘1973’]. Appendix A classification. In V. Ahmadjian & M. E. Hale (Eds), The lichens: 599-632.
New York.
Powell, C. McA., Johnson, B. D. & Veevers, J. J. 1981. The early Cretaceous breakup of eastern
RAMALINA IN AUSTRALIA Zoe
Gondwanaland, the separation of Australia and India, and their interaction with southeast Asia. In A.
Keast (Ed.), Ecological biogeography of Australia: 17-29. Den Haag.
Pryor, L. D. 1981. Australian endangered species: Eucalypts. Australian National Parks and Wildlife
Service. Special publication [5]. Canberra.
Rasanen, V. J. P. B. 1943. Das System der Flechten. Acta. bot. fenn. 33: 1-82.
— 1949. Lichenes Novi V. Soumal. eldin-ja kaswit. Seur. van. Tiedon. Poytak. 3: 178-189.
Raven, P. H. 1973. The evolution of Mediterranean floras. In F. de Castri & H. A. Mooney (Eds),
Mediterranean type ecosystems, origin and structure: 213—223. Berlin.
—— & Axelrod, D. I. 1974. Angiosperm biogeography and past continental movements. Ann. Mo. bot.
Gdn 61: 539-673.
Rogers, R. W. & Stevens, G. N. 1981. Lichens. Jn A. Keast (Ed.), Ecological biogeography of Australia:
593-603. Den Haag.
Rundel, P. W. 1978a. Evolutionary relationships in the Ramalina usnea complex. Lichenologist 10:
141-156.
— 1978b. Ecological relationships of desert fog zone lichens. Bryologist 81: 277-293.
— 1982. The role of morphology in the water relations of desert lichens. J. Hattori bot. Lab. 53: 315-320.
— & Bowler, P. A. 1974. The lichen genus Trichoramalina. Bryologist 77: 188-194.
— —— 1976. Ramalina leptocarpha and R. subleptocarpha: a fertile-sorediate species pair. Bryologist
79: 364-369.
—_— —— 1978. Niebla, a new generic name for the lichen genus Desmazieria (Ramalinaceae). Mycotaxon
4 (3): 497-499.
Schuster, R. M. 1979. On the persistence and dispersal of transantarctic Hepaticae. Can. J. Bot. 57:
2179-2225.
Sheard, J. W. 1978. The comparative ecology and distribution and within-species variation of the
lichenized Ascomycetes Ramalina cuspidata and R. siliquosa in the British Isles. Can. J. Bot. 56:
939-952.
Shirley, J. 1888. The lichen flora of Queensland, with descriptions of species. Part 1. Proc. R. Soc. Qd 5:
80-110.
Sipman, H. J. M. 1983. A monograph of the lichen family Megalosporaceae. [Bibliotheca lichenologica
18.] Vaduz.
Smith, A. L. 1921. Lichens. Cambridge.
Specht, R. L. 1970. Vegetation. In G. W. Leeper (Ed.), The Australian environment: 44-67. 4th ed.
Melbourne.
—— 1981a. Major vegetation formations in Australia. In A. Keast (Ed.), Ecological biogeography of
Australia: 163—297. Den Haag.
—— 1981b. Biogeography of halophytic angiosperms (salt-marsh, mangrove and sea-grass). In A. Keast
(Ed.), Ecological biogeography of Australia: 575-589. Den Haag.
— 1981c. Evolution of the Australian flora: some generalizations. In A. Keast (Ed.), Ecological
biogeography of Australia: 783-805. Den Haag.
Steiner, J. 1904. Flechten, auf Madeira und den Kanaren gesammelt von J. Bornmiiller in dem Jahren 1900
und 1901. Ost. bot. Z. 54 (10): 351-365.
Stevens, G. N. 1978. Lichens on mangroves along the east coast of Australia. M.Sc. thesis (unpublished),
University of Queensland.
— 1982. Ramalina leiodea (syn. R. boninensis) — a common maritime species in Oceania. Lichenologist
14: 39-45.
— 1983a. Tropical-subtropical Ramalinae in the Ramalina farinacea complex. Lichenologist 15:
213-230.
— 1983b. Clarification of the name Ramalina linearis. Lichenologist 15: 99-102.
Stizenberger, E. 1862. Beitrag zur Flechtensystematik. Ber. Tat. St Gall. naturwiss. Ges. 3: 124-182.
Tehler, A. 1983. The genera Dirina and Roccellina (Rocellaceae). Op. bot. Soc. bot. Lund. 70: 1-86.
Vainio, E. A. 1890. Etude sur la classification naturelle et la morphologie des lichens du Brésil. Pars prima
(1-29). Acta Soc. Fauna Flora fenn. 7 (1): 1-247. Pars secunda. Acta Soc. Fauna Flora fenn. 7 (2):
1-256.
Watson, W. 1929. The classification of lichens. Part 1. New Phytol. 28: 1-36.
Webb, L. J. & Tracey, J. G. 1967. An ecological guide to new planting areas and site potential for Hoop
Pine. Aust. For. 31 (3): 224-239.
Weber, W. A. & Wetmore, C. M. 1972. Catalogue of the lichens of Australia exclusive of Tasmania. Beih.
Nova Hedwigia 41: 1-137.
Wetmore, C. M. 1963. Catalogue of the lichens of Tasmania. Revue bryol. lichén. 32: 223-264.
eee
G. NELL STEVENS
Wilson, F. R. M. 1889. Australian lichenology. Proc. R. Soc. Od 6: 549-553.
Zahlbruckner, A. 1907. Lichenes (Flechten) B. Specieller Teil. In A. Engler & K. Prantl (Eds), Die
Natiirlichen Pflanzenfamilien 1: 49-249. Leipzig.
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,
153% lod 9521482217,
Bistortae 116
Bitectae 116
boninensis 207
f. subcalicariformis 207
bourgaeana 129
brevis 120, 163, 166, 214*
var. brevissima 120, 193, 194
Bulbothrix 146
caespitella 121, 123*, 129, 138, 139,
143, 149, 150, 153*, 155, 156*
calcarata 146, 186, 194
calicaris 217
f. Ecklonii 207
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
capitata 121, 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
Ciliate 116
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
Dievernia 117, 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,
25"
var. nervulosa 120, 195, 197, 200
var. squarrosa 120, 170, 172,
Z5*
fastigiata 115, 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*2161 1627:
178
fissa 131, 132, 138, 143, 149, 150,
1515152, 1975 1815) 185; 191"
193, 194, 195*, 215*
Fistularia Vainio 115, 116, 126
Fistulariae R. Howe 116
Fistulariella Bowler & Rundel 110,
115,116,175 119
Fistulosae Hue 115
flaccescens 115
fraxinea 115, 150, 203, 217
B yemensis 180, 182
f. platyna 194, 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,
PAC SERN Wel
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,
1435149; 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. fissa 120, 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*
var. 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*
Myelopoea 115, 116
Myelopoeae 116
myrioclada 120, 152, 154, 214*
nervulosa var. dumeticola 121,
122", 12575124, 134,130,138,
143, 149, 151, 196, 197, 198,
199*, 201
var. luciae 121, 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*
Niebla 115, 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*,
21S? S207
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
RAMALINA IN AUSTRALIA
pumila 189
pusilla 186, 191, 217
Ramalina 109*, 115, 149
Ramalinaceae 115, 116, 117, 118,
119
Ramalinopsis 117, 119
reductai]21--122*. 123*-129) 1132"
138, 139, 141*, 143, 149, 151,
177575176; 183
Relicina 146
rigida 158
scopulorum 115, 217
var. cuspidata 217
var. subfarinacea 217
sideriza 210
siliquosa 133, 134
sinaloensis 126
Solidae 116
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" 27.
var. leiodea 108, 127*, 136, 138,
143, 149, 152, 159, 196, 203,
205,.206% , 2072 2115.213
var. norstictica 108, 127*, 136,
138, 143, 149, 151, 196, 208,
LiL)
209* > 211,213
var. subfraxinea 125*, 127*, 136,
138, 143, 148, 149, 152, 196,
203, 204* , 207, 208, 211, 212,
213; 21687217
subgeniculata Knight 120, 185
subleptocarpha 147
subpusilla 194
tasmanica 117, 185, 215*
tayloriana 217
tenella 121, 122*, 123*, 129, 132,
136, 138, 143, 149, 151, 171,
L727 On Neel,
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*,
14351495 151156/1575 1618
17 ATT Ste Ome leh
usnea 108, 147, 154, 160, 217
Usnea 116, 118, 119, 146
Usneaceae 117, 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
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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 line illustrations, 22 maps. Hardback.
0 565 00895 1 £50.00
Titles to be published in Volume 16
Studies in the genus Hypericum L. (Guttiferae)
7. Section 29. Brathys (part 1)
By N. 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
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
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© British Museum (Natural History), 1987
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eon 8
. ‘ ge 7
ons euellCATIONS
ISSUED
27 AUG 1987
ISBN 0 565 08016 4
Botany series
ISSN 0068-2292
Vol 16 No 3 pp 225-268
a NEZ
British Museum (Natural History)*3~_
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London SW7 5BD Issued 27 August 1987
An annotated list of vascular plants collected in the
valleys south of Mt Everest
Georg Miehe ,
Department of Geography, University of Géttingen, Goldschmidtstrasse 5, D-3400
Géttingen, West Germany
Contents
ST + ype Heed period le gih ng ee eA a gy POURS eT ROME a Phe Be ey uae arts ee MeN een Tre ere Zen
REN R NN ois es cases ee tk nhs and Peto sin aieon Ce anaseace secs NT wee cuk ee teRr ay eae a Mana Tae 225
MONT ETIMEREEI STN OOE TAMRs RI css scd fo eset Sos atsa desu cinch vndschees cbnee vast natecenatecreleets 225
RPONETAMS OE CHIE IIE 65s soi sivn'c ods tides viv dhe sacgeie vaste ee kent tau hale ai ncocescan na seen cness 226
PRMEERAMNI GL CTC Clk NE MORMCUOND 60's ca taick tps bce ceniagadee hbeesrsaeesswads vndeseseevcd Sueuckaeveres 228
PEUATES On ee TE CONOR a. cil os Eas hag nasi Rs inc ee ee dig date steed tenabese ys 231
Remarks on the influence of grazing, burning,
re ee 1 Pa ARNO IR ni tye IEE PRR OE Der PERE Tas Oe te LMBE on ey ee 232
Alphabetical list of vascular plants, their life forms,
associated formations, and records of collection ..................cccceccecceeeeeeeeeeess 233
BY CE DISHES LONE BVIIDONS oc 2isc nies on cacenan ccna can dOd toe cveidedanst nen sisdencaseenetees 233
BG Uy ae TCR INN SYM oes 5c 65 con sing saicase sid Sad eeneinssacrtesesesysasstnuontecdeen 234
PROCS VIMNIOR TST OL CONGCIORS’ NAICS ii. isc: Ji cos disses soades eeetacssasaennentacinba gases deans Za
PASM aDEtUCAl TISE OL HOWETINE DIAING i cc5ics cass 9105340 c00- 7s ogee ksdosvs optete<dsapeeesndbeenasaes 255
Pupmabvoucal list of ferns and fOr BNIES. 6555. oeceuses sscssass-sanoeezes sdepugaersene tee stol 261
PACMAN TUTOR TEMES 8 uae cath nwa ne Fou Coden cgaET eg aves Cea nua teen ane den eMac aEsTS ae Te 263
References and other relevant literature ......... De OPEN ey Eee ET Pee ee aire 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° 10’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 validate 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) climatic 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° 10’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
Nang Pa la
5716 Cho Oyu 86°45’E
8153 > 'yachung Kang @ main settlement
a N me © summer settlement
—)— Cols and passes
(se 4 ) Glacier
0 1
28°N \ Mt. Everest : a
£8 ee \ ° 8848
Source:
; Khumbu Area Nepal,
Mountain Hazards Index Map;
Nepal 1:506.880 East Sheet
i Chhu
SAVeGo THY
86°45’ 87° 87°15"
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
PLANTS SOUTH OF MT EVEREST 227
Chaunrikharka (2743m) 2283,6mm
(mean monthly temperatures
of Namche Bazar +0,58°C /
100! /1949-1962)
°c Namche Bazar (3440m) ca. 6,6° 1048 mm
80 = (1948-1963, 1953?-1976) - 120
bs mm
50 100 50 - | L 100
40 4 L 80 40 - + 80
30 + + 60 30 4 + 60
20 4 L 40 20 4 + 40
> aa
10 4 L 20 10 - ~ N L 20
-7,9°
0 T T T D T T T T T T a7 0 0 al T T T T Ai T T ql a T T 7 0
Ce VE NY Wel OC exe 00 ON Oe i a i Vi VO ee OE
Months Source: Dhar & Narayanan 1965, Joshi 1981 Months Source: 0. N. Dhar, J. Narayanan 1965,
Fi 2 D P. Josh 1981
'g. Fig. 3
‘Lhajung’ (4420m) -1,27°C
(27°53,6’ N/86°49'7” E.Gr.) 528mm
(1974, 1974)
¢ mm
ne”
5° - 100
40 + + 80
30 - + 60
20 4 L 49
10 4 L 20
LU N i
-113° L :
“Mr yp on we vw we vm x x Nox
Months ho
Source: K. Higuchi (Ed.) 1984, App. 3
Fig. 4
Figs 2-4 Climatic diagrams. After Walter & Lieth (1966).
for graphical reasons, if more than 100 mm. Where the temperature line lies above the rainfall
line 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 (Firer-
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 Karnali 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 solifiuction belt is
squeezed into a narrow zone between the glaciers and the vegetation cover of the alpine belt.
Thus the upper limit 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 of 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
lichens 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 ferests 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 limit 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
lichens. 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 believed 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 fiat 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-relief. 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 solifiluction 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 mot
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 limit of forests is controlled by temperature. The rainfall is obviously not a
limiting factor as an annual precipitation of as little 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 line 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
252 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 little 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 (Firer-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 life forms (Ellenberg, 1956: 26) as completed by Ellenberg & Mueller-Dombois
(1967).
Phanerophytes (P)
Mes Pscap = large trees (S—50 m).
Mi Pscap = small trees (2-5 m).
N Pscap = dwarf trees (< 2m).
Mi Pcaesp = tall shrubs (2-5 m).
N_ Pcaesp = normal-sized shrubs (< 2 m).
N Pgram = 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 = aphyllous (Ephedra).
b = needle-leaved.
rept = reptant.
pulv = pulvinate.
fpulv = flat cushions (Silene acaulis, Gentiana acaulis).
gpulv = globose cushions (Androsace helvetica).
lsucc = 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
Hrept = creeping and/or matting.
Hscap = scapose without rosette.
Hros = rosette.
Hsem = rosette, but with leaves on the stalk.
Hscap = without rosette, mainly tall forbs.
Geophytes (G)
Survival organs protected in the soil.
Grad = 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,
Teptaces):
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.
obl 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).
+ Not included in Hara, Stearn & Williams (1978), Hara & Williams (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/11500
ft
III cloud forest (upper belt) between 3200 m/10500 ft and the upper tree line (4200 m/13 800 ft) with
Abies spectabilis, Betula utilis, and Rhododendron thickets. Beard lichens (Usnea longissima) are
characteristic.
PLANTS SOUTH OF MT EVEREST px
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/16 500 ft.
VII mostly north-facing, wind-sheltered moist alpine scrub (Rhododendron anthopogon, R. setosum,
R. nivale) between 3800 m/12 500 ft and 5200 m/17 000 ft.
VIII flushes.
IX Kobresia pygmaea — mats between 5000 m/16500 ft and 5500 m/18 000 ft.
Free solifluction belt
X phanerogams adapted to the solifluction moved scree up to the highest open slopes at about 6000
m/19700 ft.
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 of Arenaria 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.
r = 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. Schilling. 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.
WwW = T. Wraber. Cited specimens in BM and according to determination list (BM).
Z = A. Zimmermann. Cited specimens according to duplicates in BM and according to Candollea
15-17.
Alphabetical list of flowering plants
The plant names and their authors follow An 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 caudatum Wall. *
A. pectinatum Wall. ex Pax *
A. stachyophyllum Hiern
Aconitum balfourii Stapf
A. ferox Wall. ex Seringe *
A. heterophyllum Wall. ex Royle
A. hookeri Stapf *
A. laciniatum (Brihl) Stapf var.
laciniatum
A. rotundifolium Kar. & Kir.
A. spicatum (Brihl) 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
aHsem
a NP caesp
a Mi P scap
a Mi P scap
a Mi P scap
a G bulb
a G bulb
a G bulb
a G bulb
a G bulb
a G bulb
a G bulb
a Grad
a Grad
a Grad
aG rhiz
a Grad
a Grad
a Grad
a Ch suff
sobl.d EPL
a H caesp
a H caesp
a H caesp
a H caesp
a H ros
a H caesp
aH rept
a Grad
aGrad
a/e Mes P scap
a H scap/Ch suff
af Ch herb pulv
aHsem
Lagi <ahee:
Sch.2342, W.436
By.12, D.337, 380,
BL.2058, 2135, M.903,
Mc.322, Z.694, 1474,
Sm.s.n.
BL.2090, Z.446, 760
BL.2172, By.47, 326, 640,
dH.2967, Sch.2348,
2349, 2261, 2262
§.2246, 2247, 5946
Sch.2361
Z.1796
dH.2900, S.6577, P.s.n., P.
B.78, Z.1309, 1661
Z.1796 A
By.1132, H.50, M.1009,
P.s.n., $.6595, W.196,
Sm.s.n.
S.6577
P'sin.
M.875, P.s.n., W.289
Z.1889
Z.187
M.845, Z.1691A
D.404, P.s.n.
D.340, dH.1970, L.0656,
Sm.s.n., Sch.2040, Z.667
P.R.162
By.993
W.282
P.69
S.4488
P.s:n.
M.892b, W.366a
W.429
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.
M.1037, Sm.s.n.
By.169, 314, 385, 418, 742,
H.3, P.G.298
A. margaritacea (L.) Benth. *
A. royleana DC.
A. tenella DC.
A. triplinervis (Sims) C. B.
Clarke *
A.triplinervis var. intermedia
(DC.) Airy Shaw
A. triplinervis var. monocephala
(DC.) Airy Shaw *
A. xylorhiza Sch.Bip. ex Hook.f.
Andropogon munroi C. B. Clarke
Androsace delavayi Franch. *
A. geraniifolia Watt
A. globifera Duby *
A. hookeriana Klatt
A. lehmannii Wall. ex Duby *
A. nortonii Ludlow
A. sarmentosa Wall. *
Anemone demissa Hook.f. &
Thoms. *
A. obtusiloba D. Don*
A. rivularis Buch.-Ham. ex DC. *
A. rupestris Wall ex Hook.f. &
Thoms.
A. rupicola Cambess. *
Arabidopsis himalaica (Edgew.)
O. E. Schulz
A. lasiocarpa (Hook.f. &
Thoms.) O. E. Schulz
Arctium lappa L.
Arcyosperma primulifolium
(Thoms.) O. E. Schulz
Arenaria bryophylla Fernald *
A. ciliolata Edgew.
A. debilis Hook.f. ex Edgew. &
Hook.f.
A. densissima Wall. ex Edgew. &
Hook.f. *
A.edgeworthiana Majumdar
A. festucoides Benth.
A. glanduligera Edgew. ex
Edgew. & Hook.f. *
A. glanduligera var. cernuaN. F.
Wilson
A. globiflora Edgew. & Hook.f.
A. melandryoides Edgew. ex
Edgew. & Hook.f.
A. polytrichoides Edgew. ex
Edgew. & Hook.f.
A. pulvinata Edgew. ex Edgew.
& Hook.f.
aH sem
a H rept/sem
aH rept/sem
aHsem
aHsem
af Ch herb pulv/a
H sem
af Ch herb pulv
a H caesp
e f Ch herb pulv
a H ros
e f Ch herb pulv
aH ros
e g Ch herb pulv
a Hros
a Hros
aHsem
aHsem
aHsem
aG bulb
a G rhiz
Tsem
T caesp
a H caesp
aH ros
a g Ch herb pulv
a H rept/ros
a H caesp
a g Ch herb pulv
a g Ch herb pulv
a g Ch herb pulv
a f Ch herb pulv
af Ch herb pulv
a g Ch herb pulv
a Grad/rept
a g Ch herb pulv
af Ch herb pulv
PLANTS SOUTH OF MT EVEREST
II
II
Vv
Vv
Vv
VI
VI, VI, IX
IV
IX, X
je 4
IX
Vill
IX
IX
b, SE Vi evil,
VIII
iil, V, x
By,
Vv
VII
237
By.370, dH.2879
W.303
D.384, dH.2917, Z.1755
By.411, 1130, D.332,
Z.1089
£167; 72.8.0;
D.343, M.1015, Sm.s.n.
H.47, BL.2110, D.414,
W.210, Sm.s.n.
M.817b, Z.1733
BL.2123, Sm.s.n.
S.4592
D.416, Mc.323, M.1045,
$.637
Z.823
BL.2008, D.390, Mc.284,
Z.816
BL.2046, By.445, D.412,
Mc.318, Sm.s.n.
By.46, 227, 1092, dH.2923,
Z.466, 686
BL.2109, D.423, Z.601,
693, Sm.s.n.
P.R.306
By.393, M.902, P.s.n.
BL.2087
S.669
By.121, W.297
B.&Sh. 5656
f.M.
Mc.52
£1.30; M10/6, P:s.n.., f.
Heub.
H.41, M.1043, Sm.s.n.
By.637
By.797, D.410, Mc.2038,
S:7115
Mc.377, W.227, Sm.s.n.
Sw.s.n.
By.385, H.28, M.955,
Mc.377, P.s.n., W.178,
231, Sm.s.n.
M.1074
BL.2073, Mc.309, S.7095
W.189, Sm.s.n.
M.989, S.7157
By.773, Sw.s.n.
238
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. stricta Edgew.
A. wallichiana Besser *
Aruncus dioicus (Walter) Fernald *
Arundinaria maling 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 Grah. ex Benth.
Begonia picta Smith *
Berberis angulosa Wall. ex
Hook.f. & Thoms. *
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.
& Thoms.) Engl.
Betula alnoides Buch.-Ham. ex.
D. Don *
B. utilis D. Don *
Bistorta affinis (D. Don) Greene
*
B. amplexicaulis (D. Don)
Greene *
G. MIEHE
a G bulb
aG bulb
a G bulb
aH scap
a H scap
aH scap
a H scap
aH scap
a H scap
a Ch suff
a/e NP gram
a H sem/Ch suff
a G rhiz/sem
aHsem
a G rhiz/sem
a G rhiz
a H sem/Ch suff
a G rhiz/rept
a H sem/Ch suff
aGrad
a H scap
a H rept/caesp
aH rept
a H scap
a H caesp/rept
s Ch frut rept
a/e G bulb fac E
a NP caesp
a/e NP caesp
a/s NP caesp
a/e NP caesp
a/e NP caesp
a Ch frut
a NP caesp
a/e NP caesp
a G rhiz/ros
a Mes P scap
a Mes P scap
a G rhiz
a Grad
BL.2064
BL.2029
BL.2055
L.0181, P.G.268
Z.1792
Z.1655
P.s.n.
M.836, P.s.n., W.200,
Z.1792
B.5707, dH.2997, S.6589
By.297, 464, Beer 25447
W.347
M.828, Mc.392, W.361,
Z.1635, 1653
D.386, 415
By.696, dH.2953a, M.899,
P.R.298, Z.1717, 1822,
Sm.s.n.
BL.2128, D.415, Sm.s.n.
By.1169, M.869, 1042,
Sch.2173
Sch.2210, 2250, Z.1880,
1885
Beer 25388, By.564,
Mc.313
By.321, Z.1768A
BE.2155;S8:7100
BL.2132, By.210, 311, 312,
339
S.7ko2
P.s.0.
M.1013, Mc.314
P77.
Sch.2270, Z.494, 1619
1154, Z.1758
BL.2103
Z.593
Z.480
By.175, Sch.2323, Z.653
f.M.
By.1157, W.434, Z.406,
492
By.478, D.365, Z.1481,
Z.1573, Sm.s.n.
By.153, 211, M.849, 913,
P.G. 348, Sch.2344
B. macrophylla D. Don) Sojak *
B. milletii Lév.
B. perpusilla (Hook.f.) Greene
B. vaccinifolia (Wall. ex Meisn.)
Greene *
B. vivipara (L.) S. F. Gray
Biswarea tonglensis (C. B.
Clarke) Cogn.
Boschniaka himalaica Hook. &
Thoms. ex Hook.f. *
Brachyactis anomala (DC.)
Kitam.
Braya oxycarpa Hook.f. &
Thoms. *
Buddleja crispa Benth. *
Bulbophyllum retusiusculum
Reichenb.f.
Bupleurum candollii Wall. ex
De:*
Cacalia pentaloba Hand.-Mazz.
Calamagrostis emodensis Griseb.
C. epigejos (L.) Roth
C. pseudophragmites (Hall.f.)
Koel. ex. Baumg.
Calanthe alpina Hook.f. ex Lindl.
Callitriche palustris L.
Campanula aristata Wall.
C. modesta Hook.f. & Thoms. *
C. nakaoi Kitam.
C. pallida Wall.
Capsella bursa-pastoris (L.)
Medikus
Cardamine inayatii O. E. Schulz
a
C. loxostemonoides O. E. Schulz
*
C. scutata subsp. flexuosa
(Withering) Hara
C. trifoliolata Hook.f. & Thoms.
Cardiocrinum giganteum (Wall.)
Makino *
Carex atrata subsp. pullata
(Boott) Kiikenthal
C. atrofusca Schkuhr
C. inanis Kunth
C. infuscata Nees
C. laeta Boott
C. lehmannii Drejer
C. munda Boott
C. nakaoana T: Koyama
PLANTS SOUTH OF MT EVEREST
a G rad/sem
a Grad
a G rad/rhiz
a Ch frut rept
a G rad/sem
st TL
aG bulb
aHsem
a G rad/H caesp/f
Ch herb pulv
a NP caesp
obl e E Ch succ
aGrad
aH scap
a H caesp
a H caesp
a H caesp
aGrad
a hyd H caesp
aHsem
aH scap
a G rhiz
aH scap
T sem
T caesp
a H/T scap
T scap
T scap
a G bulb
a G rhiz
a G rhiz
a G rhiz/caesp
a G rhiz
a H caesp
a G rhiz
a G rhiz/caesp
a H caesp
VI, VII, 1X
Vv
VI, IX
Ill
VI, VII, 1X
I
Vv
Vv
X
VIII
IV, VI, VII
239
BL.2198, M.948, Mc.330,
P.s.n.,$.4521, W.201,
210 Z3ta1s 1418,
Sm.s.n.
W.s.n., Z.1478
Beer 9552
By.75, 82, 137, 174, 348,
502, M.861, Sch.2346,
2175, 2181, Z.609, 1709
D.366, 391, Z.640, Sm.s.n.
§.7190, Z.785
Mc.306
M.911
Z.564, 597
Z.262
L.0433
| BRC Re Pe 2
B.5651, M.978
W.394
ZATTY
Sm.s.n.
Mc.265
M1114, 2 s:n:
By.427, D.462, Mc.361
By.549, 581, L.2138,
Sm.s.n.
S.1666, W.295
By.307, 315, Z.1654
By.214, P.78, Z.418, 1764
BL.2016
BL.2111, Mc.360
Z.788
Mc.272
P.s.n., Sch.2248, Z.1898
BL.2166, D.446
D.376, M.1094, P.G.288,
R.331
BL.2160
M.958, 1014
P.R.338, Z.696
P.R-EZ1 R325
W.310
dH.2994, M.1029, P.
G219,; 6.29230 .297,
S.666, W.169
240
Carpesium nepalense Less.
Cassiope fastigiata (Wall.) D.
Don *
Castanopsis tribuloides (Sm.) A.
DC.
Celastrus stylosus Wall.
Cerastium fontanum Baumg.
Chamaesium novemjugum (C. B.
Clarke) C. Norman
Chenopodium botrys L.
C. graveolens Willd. +
Chesneya nubigena (D. Don) Ali
*
Chionocharis hookeri (C. B.
Clarke) J. M. Johnston *
Chrysosplenium alternifolium
var. sibiricum Seringe ex DC.
C. carnosum Hook.f. & Thoms. *
C. cf. griffithii Hook.f. & Thoms.
C. lanuginosum Hook.f. &
Thoms.
C. nudicaule var. intermedium
Hara
Cicerbita cyanea (D. Don)
Beauv.
C. macrantha (C. B. Clarke)
Beauv. *
C. macrorhiza (Royle) Beauv.
C. macrorhiza var. saxatilis
(Edgew.) Beauv.
Circaea alpina subsp. micrantha
(Skvortsov) Bufford +
C. alpina subsp. imaicola
(Aschers. & Magnus) Kitam.
C. repens Wall. ex Aschers. &
Magnus
Cirsium falconeri (Hook.f.)
Petrak *
C. verutum (D. Don) Spreng. *
Clematis montana Buch.-Ham. ex
DC:>
Clinopodium umbrosum (M.
Bieb.) C. Koch
Clintonia udensis var. alpina
(Kunth ex Baker) Hara *
Coccinea grandis (L.) Voigt
Codonopsis ovata Benth. *
C. thalictrifolia Wall.
*
Coleus barbatus (Andrews)
Benth. *
Colutea multiflora Shap. *
Commelina paludosa Blume *
Corallodiscus lanuginosus (Wall.
ex DC.) Burtt *
Cortia depressa (D. Don) C.
Norman *
G. MIEHE
a H scap
s Ch frut
s Mes P scap
ad LP frut
aH rept
a Grad
T scap
a f Ch frut pulv
e g Ch herb pulv
H caesp/scap
H caesp
a G rhiz/scap
a G rhiz/sem
G rhiz/scap
a Grad/H scap
a Grad/H scap
a Grad
a G rad/a H scap
T rept/scap
T rept/scap
T rept/scap
a G rad/sem
a G rad/scap
ael PL frut
a H caesp
a G rhiz
owe
a H scap
a G rad/rept
a H scap/Ch suff
a NP caesp
aH scap
e Hros
aGrad
II
W.397
By.469, 687, D.379,
W.248, Z.612, 634, 702,
1559, 1756, Sm.s.n.
f.M.
S.4484
By.498
M.936
By.724, P.s.n.
By.1124
BL.2112, By.41, M.1035,
Mc.329, W.249,
Sm.s.n.
Sw.224, 225
Z.805
By.38, S.7117
M.983
Z.458
BL.2035, Mc.53
dH.2950a, P.s.n., Z.1791
P:s.n.
By.586, 712, P.s.n.
dH.2941, M.128
W.317
P:7
S.5750
W.377, Z.1670
By.601
By.654, dH.2975,
Sch.2186, Z.502
P.s.n., W.307, Z.1670
By.26, Z.641
Psa.
By.362
By.342, 430, 453, M.1058,
Sm.s.n.
Z.1833, 1865
dH.2951, Sch.2165
By.465
D.321, Mc.394, M.811,
Z.422, 1225
BL.2179, D.368, P.s.n.,
G.259, Sm.s.n.
Cortiella hookeri (C. B. Clarke)
C. Norman *
Corydalis cashmeriana Royle *
C. casimiriana Prain
C. cavei D. G. Long
C. chaerophylla DC.
C. chasmophila Ludlow
C. flabellata Edgew.
C. flaccida Hook. f.& Thoms.
C. govaniana Wall. *
C. hookeri Prain
C. juncea Wall. *
C. leptocarpa Hook.f. & Thoms.
C. longipes DC.
C. meifolia Wall. var. meifolia *
C. polygalina Hook.f. & Thoms.
C. stracheyi Duthie ex Prain +
Corylus ferox Wall. *
Cotoneaster acuminatus Lindl.
C. adpressus Bois
C. frigidus Wall. ex Lindl. *
C. integrifolius (Roxb.) Klotz
C. meuselii Klotz
C. microphyllus Wall. ex Lindl. *
C. nitidus Jacques
C. sanguineus Yi
Cremanthodium decaisnei C. B.
Clarke *
C. ellisii (Hook.f.) Kitam. *
C. nepalense Kitam. *
C. oblongatum C. B. Clarke *
C. reniforme (DC.) Benth. *
C. retusum (Wall. ex Hook.f.) R.
Good *
Crepis tibetica Babcock
Cryptothladia polyphylla (Wall.
ex DC.) 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
Cyanotis vaga (Lour.) J. A. & J.
H. Schultes *
Cymbidium hookerianum
Reichenb. f. *
PLANTS SOUTH OF MT EVEREST
aGrad IX, X
a G bulb V, VII
Vv
Vv
I
a H caesp VII
t
Ill
a H caesp III, V
VII
a G bulb/scap LEY
Ill
a G rad/caesp Ill
a H caesp Vil
a G bulb vil
a H caesp VI
a Mes P scap I
a N P caesp He
a Ch frut rept II, V
a Mi P caesp II
s Ch frut rept iV
a N P caesp II
s Ch frut rept Me
s/a N P caesp II, V
a N P caesp II
a Hros VIII, X
aHros VIII, X
aH ros VII
aHros VII, X
aHsem vil
aH sem VII
a H ros/sem ?
aGrad Vv
ELT Par II
aH rept V, VI
a Grad/H rept Vi¥i
a H rept/scap Vv
a G rad/H rept Vv
a G rad/H rept Vv
aH rept V, VI
a Grad/rept VI, VII
aH II
fac. e Ch E succ I
241
M.1001, W.164, 233,
Sm.s.n.
D.431, Mc.283, P.s.n.
H.10 (f. Lidén)
H.10 (f. Lidén)
Mc.266
BL.2163, $.628
By.193
Mc.295, S.8360
M.879, Z.809, Sm.s.n.
S.7153
BL.2164, Mc.341, $.4538
S.4536
By.525, 1167, dH.2935,
M.850, W.386
M.1083, Beer 9555,
Sm.s.n.
Mc.327
1:357
S.4606
dH.2930, M.854, Mc.273,
S.4614, Sch.2030
Sch.2031, 2263
Sch.2442A
9.7125
Sch.2355
By.1089, 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.A7,.33,,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 L. *
Delphinium caeruleum Jacquem.
ex Cambess.
D. drepanocentrum (Brihl) Munz
*
D. glaciale Hook.f. & Thoms.
D. cf. nepalense Kitam. &
Tamura
D. viscosum 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 (Lout.)
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
ad PL frut
a H scap
a H scap/caesp
a G rhiz
a Grad
a H caesp
a H caesp
s NP caesp
s NP caesp
a H scap
aH sem
aHsem
aHsem
aHsem
aH sem
a NP caesp
a NP caesp
a NP caesp
a NP caesp
a NP caesp
a G rhiz
ad HL
aHsem
aHsem
a G bulb/scap
a G bulb/scap
a G bulb/scap
a Grad
s Ch frut/rept
a H sem/scap
a G rad/scap
a Grad
s Mes P scap
a H ros/a f Ch herb
pulv
aHsem
aHsem
—
a
II, V-VIII
II
I, VI
Mc.395
B.5659, Mc.275, P.193
By.397, 872
BL.2199, By.120, D.395,
Sm.s.n.
By.142
M.892c, P.s.n.
M.877, 851b, W.349
S.4480
W.328
f.M.
P:s.n;
By.423, 537, 621, dH.2969,
L.0191, M.882
M.1060, P.s.n., W.172,
219, Sm.s.n.
M.1033
Beer 25387, W.372
Sw.71-72
By.364
Pes hi:
Mc.38, $.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
Z.749B
Z.749
Z.746
S.7163
S.7164
L.0529
Beer 25361, Sw.441
Z.1671
Z.765
Sch.2366A
S.6559
Mc.325, W.167
BL.2178, D.424. S.7144
BL.2114, D.440, Mc.287,
Z.521, 581
D. ellipsoidea Hook.f. & Thoms.
D. gracillima Hook.f. & Thoms.
D. lasiophylla Royle
D. lasiophylla var. leiocarpa
(Pamp.) O. E. Schulz
D. cf. oariocarpa O. E. Schulz
D. oreades Schrenk *
Drosera peltata Smith
Drymaria cordata (L.) Willd. ex
Roem. & Schult.
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 Raven
Erigeron bellidioides
(Buch.-Ham. ex D. Don)
Benth. ex C. B. Clarke *
E. kumaunensis (Vieth.)
Wendelbo
E. multiradiatus (Lindl. ex DC.)
C. B. Clarke *
Eriophyton wallichii Benth. *
Erysimum hieraciifolium L.
Euonymus amygdalifolius
Franch.
E. frigidus Wall.
E. hamiltonianus Wall. *
E. pendulus Wall.
E. porphyreus Loesener +
PLANTS SOUTH OF MT EVEREST
T caesp Vv
T sem V, VII
af Ch herb pulv/a IX, X
H ros
a f Ch herb pulv VI, VII
a Hros/a f Ch herb VII, IX
pulv
aHros/afChherb X
pulv/G rhiz
aH ros III, V
aHrept/afChherb IX,X
pulv
a H scap/G rad V, VI
a/e Mes/Mi P I
scap/caesp
T scap/caesp TV.
a H scap Vv
T scap | | "3
T scap 8 hf
aNP caesp/Ch suff iI
T scap II, Ill
a H caesp ViVi
aH caesp VI
s/a Mi P caesp if
p Ch frut LEV VIS
aH scap V, VI
a H scap ?
a H scap IV, VII
?
aHsem II
aHsem I, V
aHsem V, VI
a G rhiz/scap X
T scap II, V
s NP caesp I
s NP caesp I
a Mi P scap |
s Mes P scap I
a Mi P scap I
243
W.204
Z.807
BL.2075, M.1102, Sm.s.n.,
S.7158, W.175, 184
D.445
P.G.206
BL.2190, M.1098, Z.569
By.365, D.333, Z.1694
BL.2119
By.583, 590, 591, dH.2962,
M.880, P.s.n., W.330,
Z.8:n.
P.s.n., Sch.2442A, S.6574,
Z.1850
Z.1778
Z.683
By.205, 675, D.339,
L.0180, M.889
Z.643
By.509, 611, 882, 1123,
M.822, P.s.n., Z.1654,
1828
M.889a, P.s.n., W.302,
£31120
M.851la
P:s-n:
BL.2013, Sch.2336, Z.426
By.736, BL.2069, D.439,
M.993, P.R.333, G.293,
Sch.2102, W.258,
Sm.s.n.
By.310, P.R.336
W.384, 392
M.844
W.300
D.325, Z.447, 451, 727
Z.1864
By.294, M.876, Z.451,
1565, 1622
By.1173, M.1007, W.236,
2.1510; Sm.s.n;
Z.450
S.6573
P.8.7:
By.988
$.6572
Mc.43, $.5949
244
E. tingens Wall. *
Euphorbia himalayensis Klotzsch
E. longifolia D. Don
E. pseudosikkimensis (Hurusawa
& Ya. Tanaka) Radcliffe-Smith
E. sikkimensis Boiss.
E. stracheyi Boiss.
E. wallichii Hook.f. *
Euphrasia himalayica Wettst. *
E. platyphylla Pennell
E. secundiflora Pennell
Eurya acuminata DC. *
Fagopyrum dibotrys (D. Don)
Hara
F. esculentum Moench
Festuca ovina L.
F. polycolea Stapf
Festuca tibetica (Stapf) Alexeev +
Ficus sarmentosa Buch.-Ham. ex
Sm.
Fragaria nubicola Lindl. ex
Lacaita *
Fritillaria cirrhosa D. Don
Galearis stracheyi (Hook.f.) P. F.
Hunt *
Galinsoga parviflora Cav.
Galium aparine L. var. aparine
G. asperifolium Wall.
G. asperifolium var. sikkimense
(Gandoger) Cufodontis
G. asperuloides subsp.
hoffmeisterii (Klotzsch) Hara
Gaultheria fragrantissima Wall. *
G. hookeri C. B. Clarke
G. nummularioides D. Don
G. pyroloides Hook.f. & Thoms.
ex Mia. *
G. trichophylla Royle *
Gentiana algida Pall. *
G. algida var. nubigena (Edgew.)
Kusn.
G. algida var. parviflora (C. B.
Clarke) Kusn.
G. algida subsp. przewalskii
(Maxim.) Kusn.
G. argentea (D. Don) C. B.
Clarke
G. capitata Buch.-Ham. ex D.
Don *
G. crassuloides Bur. & Franch.
G. depressa D. Don *
G. MIEHE
s Mi P scap
a G rad/scap
a G rad/scap
a G rad/scap
a G rad/scap
a Grad/rept
a G rad/scap
T scap
T scap
T scap
s Mes P scap
aHsem
a H scap
a H caesp
a H caesp
a H caesp
sr PL frut
a H caesp
a G bulb
a G rhiz
T scap
T rept
T rept
T rept
s NP caesp
s NP caesp
s Ch frut rept
s Ch frut rept
s Ch frut rept
e H caesp/rept
e H rept/sem
e Hrept
e H rept/sem
e H scap
e H ros
e H caesp
e f Ch herb pulv
TV val
VI, VI
Vv
Ill, V
Ill
IV
Ill, V
S.4607
Sch.2086, 2291, 2324
Sch.2069
dH.2959
Sch.2060
B.5698, BL.2131, M.954,
Z.490, 410, 537, Sm.s.n.
By.101, 1127, M.826, Z.378
By.415, Sm.s.n.
S.656
P.s.n.
S.4482
Z.766, 1862
By.304, 701
Mc.289, W.183, 242,
Sm.s.n.
D.373, $.550
M.1072
f.M.
By.1129
Z.690, Sm.s.n.
By.131, D.427
Z.1765
By.636, P.s.n.
W.426
P-R275
W.306
Sch.2050, Z.444, 1856
BL.2022, S.4612
M.820
S.4616, Z.792
By.109, M.970, S.4541,
Z.623, 1785
$.6591
M.1075, P.G.203,
Sch.2313, Sm.s.n.
Sm.s.n.
H.49
E. Onyon 5
Sw.525
Sm.s.n.
By.362, H.56, M.1117,
P.147, 149, Sch.2316,
Tilman s.n.
G. elwesii C. B. Clarke
G. infelix 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. stellata Turrill +
G. strobilacea H. Sm.
G. tubiflora (G. Don) Griseb. *
G. urnula 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. & 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 ellipticaD. Don *
Hedera nepalensis K. Koch
Hedychium densiflorum Wall.
H. spicatum Smith *
Hedysarum sikkimense Benth. ex
Baker *
PLANTS SOUTH OF MT EVEREST
aH sem
a H caesp
e H sem/caesp
e H sem/caesp
e H caesp/sem/e f
Ch herb pulv
e f Ch herb pulv
H caesp
a H sem/G rhiz
e H caesp/rept
a H scap
aH rept
a H sem/caesp
e Hros
e Hros
e H ros/G rhiz
T caesp
aHros
T ros/sem
a H ros/sem
T sem/scap
T caesp
aHsem
aHros
aHsem
aHsem
aH sem/G rad
a G rad/H ros
a H scap
a G rad/rhiz
a G rhiz
a G rad/H caesp
a Grad
a H rept/a f Ch herb
pulv
a G bulb
aH scap
Tsem
sr PL frut
a G bulb
aG bulb
aH rept
VII
VIII
V, VI, VII
Vv
V, VI
VII
IX, X
X
V, VII, 1X
X
Visi
?
IX
VI, VII
VII
Ill, V
245
Beer 25368
BLM 9548
H.2, 25, M.940, P.R.291
dH.2963, Sm.s.n.
By.597, H.35, M872,
P.R.290, G.212, S.1673,
6587, Sch.2314, Sharma
E.465, W.220, Sm.s.n.
M.1082
Sw.428
S.589
Sch.2184
W.158
P.B.9, 153, 154, Sch.2081
D.2996, P.s.n.
P.G.287, S.1674, 1694,
W.239
Sw.314
BLM 9549
M.1080, Sch.2314, $.6595,
Sw.21-23, Sm.s.n.
H.24, M.1088, W.205
M.1088
M.999, P.R.334, G.253,
G.302, Sm.s.n.
W.427
M.1110, P.s.n., W.221
S;1677
By.14, 152, D.335, 401,
M.947, S.7150, Sm.s.n.
By.1146, D.360, Sm.s.n.
By.275, P.G.329
By.192, 234, 463, BL.2096,
D.420, H.37, Mc.347,
Sm.s.n.
Z.1872
By.670, 787, P.s.n., W.353
Z.1874
M.925, W.316
By.642
By.268, BL.2065, D.345,
H.32, Mc.321, Sm.s.n.
L.0193
By.316, BL.2165, M.979,
Mc.344
H.70
BL.274
By.239, 431, M.918, W.294
Z.1838, 1840
Sch.2042
Sch.2043
BL.2079, By.43, 438,
M.1024, Mc.310, $.7101,
Sm.s.n.
246
Helictotrichon sp.
Hemiphragma heterophyllum
Wall. *
Heracleum brunonis (DC.) C. B.
Clarke
H. candicans Wall. ex DC.
H. nepalense D. Don
H. wallichii DC.
Herminium josephii Reichenb.f.
H. lanceum (Thunb.) Vuijk *
H. macrophyllum (D. Don)
Dandy
H. pugioniforme Lindl. ex Hook.f.
Hierochloe flexuosa Hook.f.
Hippophae tibetana Schlecht. *
Holboellia latifolia Wall.
H. latifolia var. angustifolia
(Wall.) Hook.f. & Thoms. *
Hydrangea heteromalla D. Don
Hypecoum leptocarpum Hook.f.
& Thoms.
Hypericum choisianum Wall. ex
N. Robson *
H. himalaicum N. Robson
Ilex dipyrena Wall. *
I. intricata Hook.f.
I. odorata Buch.-Ham. ex D. Don
I. sikkimensis Kurz
Impatiens falcifer Hook.f.
I. glandulifera Royle *
I. puberula DC.
I. pulchra Hook.f. & Thoms.
I. sulcata Wall. *
I. urticifolia Wall. *
Indigofera bracteata Grah. ex
Baker
I. heterantha Wall. ex Brandis *
Inula cappa (Buch.-Ham. ex D.
Don) DC. *
I. hookeri C. B. Clarke *
I. nervosa Wall. ex DC.
Tris kemaonensis D. Don ex
Royle *
Jasminum humile L. *
J. humile forma pubigerum (D.
Don) Grohmann
J. humile var. revolutum (Sims)
Stokes
Juncus benghalensis Kunth
J. clarkei Buchenau +
J. concinnus D. Don
J. grisebachii Buchenau
J. himalensis Klotzsch *
J. leschenaultii Gay
G. MIEHE
aH caesp
aH rept
a Grad
a Grad
a Grad
aGrad
aG bulb
aG bulb
aG bulb
a G bulb
a H caesp
a/e Ch frut/pulv
ael PL frut
ael PL frut
a Ch frut/r PL frut
a H caesp/rept
a H caesp
a H scap
s Mi P scap
s NP caesp
s MiP scap
a MiP scap
a H caesp
T scap
a H scap
aH scap
aH scap
aH scap
a H caesp
a NP caesp
a/e Ch suff
aH scap
a/e G rhiz/H scap
a G rhiz
sd PL frut
sd PL frut
sd PL frut
a G rhiz
a G rhiz
a G rhiz
a G rhiz
a G rhiz
a H caesp
M.892
dH.2978, Z.1819
By.322, 345
BL.2150, By.850, S.6598
M.883b
By.809, M.883c
D.393
H.65a
BL.2174, M.945, Mc.311
Mc.355
By.520
By.428, dH.2989, f.M.,
P.G.305, f.Sch.(2268),
2179
$.4490
Z.432, 473
Mc.269
By.768, Sm.s.n.
Mc.399, S.7168
Mc.404
Z.1675
P.73, S.4611
S.4610
BL.2019
By.246, 376
L.067
By.300, 573-575, W.341
f.M.
dH.2916, P.s.n., $.6570,
Z.1693, 1845A
dH.2919
By.81
dH.2913, Z.666
By.860
P.G.203
P.R.327
W.322
M.868, P.B.55, 171
W.376
M.1029a, $.567
Z.1724
J. leucanthus Royle ex D. Don *
J. leucomelas Royle ex D. Don *
J. pseudocastaneus (Lingelsh.)
Samuelsson
J. sphacelatus Decaisne
J. 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) Kiikenthal
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. &
Thoms. *
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
a H caesp
a G rhiz
a G rhiz
a G rhiz
b N/Mi P scap/Ch
frut
b N/Mi P scap
b Ch frut/rept
a H caesp
a H caesp
a H caesp
a H caesp
a H caesp
a H caesp
a H caesp
a H caesp
a H caesp
T rept
T rept
T rept
a H sem/G rad
aHros
aH scap
aH ros
T caesp
T caesp
aHsem
aHros
aH ros/G rad
aH sem/f Ch herb
pulv
G rhiz/ros/sem
af Ch herb pulv
IV
VIII
?
VIII, X
Ill, IV
Ill, V, VI
Ill, V
VI, VIII, 1X,
Xx
IX
VIII, X
IV, VI, VII, X
VI
IV
VIII
VIII
Ill, VI, VI
VIII, X
II
VI
VI, IX
VI, IX
~
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.1134, D.459, P.G.300,
318, $.4513, 4515, 4528,
7116
By.80, P.3, 4, $.4529, 4514,
W.410
By.1168, Z.1699
BL.2127
M.867, P.132
P.O .267
W.269
BL.2188
M.s.n., 959, 988, 1071a
M-95751071b, P.R.332,
W.237
M.1053, 1104, 1111
M.866, 1087, Mc.2032,
P.133, 134, R226
BL.2197
M.916, P.s.n., W.382
W.245, Z.651
M.981, Mc.312, W.381
M.982, 1023, W.251,
Z1529-Smisay
BL.2105, By.392, D.407,
M.1069, Sm.s.n.
L.0139
BL.2117, By.13, D.442,
Sm.s.n.
By. 668
W.208
Beer 25543
Z.403
dH.2945, Z.1754
M.881, 942, W.403
D.356, Z.1607, 1697,
Sm.s.n.
BL.2139, D.428, H.48,
M.1048, 1068, 1092,
PG.207,G.207,.8:6582;
W.166, Z.1591, Sm.s.n.
248
L. nanum (Hook.f. & Thoms. ex
C. B. Clarke) Hand.-Mazz.
L. stracheyi (Hook.f.) C. B.
Clarke ex Hemsley
Lepidium capitatum Hook.f. &
Thoms.
Leptodermis kumanoensis Parker
*
L. stapfiana H. Winkler
Leucosceptrum canum Sm. *
Leycesteria formosa Wall. *
Lignariella hobsonii (Pearson)
Baehni *
L. hobsonii subsp. serpens (W.
W. Smith) Hara
Ligularia amplexicaulis DC. *
L. fischerii (Ledeb.) Turcz. *
Lilium nanum Klotzsch *
L. nepalense D. Don *
Lindera 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.
Lloydia flavonutans Hara
L. longiscapa Hook *
L. serotina (L.) Reichenb. *
L. serotina var. parva (Marquand
& Shaw) Hara
Lobelia seguinii H. Lév. & 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. hispida Pall. ex Willd. *
L. hispida var. setosa Hook.f. &
Thoms.
L. lanceolata Wall.
L. litangensis Batalin
L. myrtillus Hook.f. & Thoms. *
L. obovata Royle ex Hook.f. &
Thoms. *
L. rupicola Hook.f. & Thoms. *
Luzula sp.
G. MIEHE
af Ch herb pulv
a H caesp
T rept
a NP caesp
a NP caesp
a NP caesp
a NP caesp
T rept
T rept
aH sem
aH scap
a G bulb
aG bulb
s Mes P scap
a/s Mes P scap
a/s Mes P scap
aGrad
a Mes P scap
s Mes P scap
aG bulb
a G bulb
aG bulb
aG bulb
a H scap
a H caesp
aHsem
aH sem
aHsem
aHsem
a Ch frut
a Mi P caesp
a Ch frut
a NP caesp
a Ch frut
a NP caesp
a Ch frut
a Ch frut
a Ch frut
a Ch/NP caesp
H caesp
IX
Vv
Vv
VI, VII
VI
II
VIII, 1X
VI-IX
BL.2125, M.1041
By.389, dH.2953, M.833,
P.111, 112, Sch.2176,
Z.1651, 1821
M.1116
S.7160
S.4618
Z.1858
P.96, Z.743
W.331
BL.2207
By.171
By.286, L.0182
BL.2056, 2074, By.150,
S.4775
By.176, 179, Sch.2248
S.6566
S.6571, 6599
Z..260
W.339
S.6567
$.5942
Mc.279
Beer 25375, Sm.s.n.
BL.2962, By.308, D.375
BL.2170, M.944, Mc.326
Z.1863
H.34, P.R.347
By.1174, M.1056, P.G.223,
R.329, $.6593, W.250
M.946, W.402
W.292, 400
W.420
Mc.393
BL.2083, 2152, By.68, 873,
M.935, Z.429
Mc.305
D.434, Sm.s.n.
BL.2104, M.1050, Mc.350
BL.2023, By.51, 282, 557,
1002, D.452, M.934
Z.520
By.425, D.418
BL.2185, By.903, P.R.357,
Sch.2078, Z.614, Sm.s.n.
By.37, M.985, P.R.358,
Z.509, 1530, Sm.s.n.
P.R.341
Lyonia ovalifolia (Wall.) Drude *
L. villosa (Hook.f.) Hand.-Mazz.
*
Lysimachia prolifera Klatt
Malaxis cylindrostachya (Lindl.)
Kuntze
M. muscifera (Lindl.) Kuntze
Mandragora caulescens subsp.
flavida Grierson & Long *
Meconopsis bella Prain *
M. horridula Hook.f. & Thoms. *
M. paniculata Prain *
M. simplicifolia (D. Don) Walp *
M. sinuata Prain *
Megacodon stylophorus (C. B.
Clarke) H. Sm. *
Meliosma dilleniifolia (Wall. ex
Wight & Arn.) 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. & Thoms.) 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 rubifloraD. 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
a/s N/Mi P scap
a H caesp/rept
a G bulb
aG bulb
a Grad/ros
a H scap
aHsem
aH sem
a H scap
a H scap
a H scap
a Mi P scap
s Mes P scap
T ros
a H caesp
aHsem
a H caesp/scap
a H scap
a H caesp
s Ch frut
s Ch frut rept
a Hscap/G rad
aH sem
a NP caesp
s MiP scap
a G bulb
a G bulb
a H scap
a H rept/scap
a Grad
PLANTS SOUTH OF MT EVEREST
III
Ill
I, Il
4 ef
V>VEIX
II
i,
i; V, Vil
TV
Vv
Vv
Ill, V—-VII
249
f.M.
M.858, P.G.358, Z.1852
Z.781
Mc.397
By.332, H.15
Z.688
Beer 25425, BL.2161
By:451,54:51,.P:s-n..
Sch.2177, 2351, W.280,
Sm.s.n.
S.649, W.395
By.28, Mc.348, Z.594
Beer 25445
BL.2153; By.271, L.0183,
Z.1703
S.4620
S.4478
M.s.n.
Z.400
Z.258
BL.2156
BL.2183, D.367, M.887,
Sm.s.n.
Sch.2063, Z.1859
Z.510
By.350, dH.2971, M.969,
P.s.n., Sm.s.n., W.432
By.220, Einarsson 9,
L.0178, M.870, Mc.444,
P YSN). cs. 3000, 1150.
Z.635, 687
L.0212
BL.2081
S.4482
H.26, M.963, P.16
H.61, P.16
B.5778
B.5636, By.52, D.385,
H.16, L.0170, M.905,
Mc.359, P.s.n., Z.1766
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 subsp. oederi var.
heteroglossa Prain *
P. pseudoregeliana Tsoong
P. punctata Decr. +
P. rhinanthoides Schrenk.
P. roylei Maxim.
P. scullyana Prain ex Maxim. *
P. siphonanta D. Don
P. trichoglossa Hook.f. *
P. wallichii Bunge
Pegaeophyton minutum Hara
P. scapiflorum (Hook.f. &
Thoms.) Marg. & Shaw *
Pentapanax leschenaultii (DC.)
Seem. *
G. MIEHE
a G rhiz
aG bulb
aG bulb
aHros
aH rept
aH rept
aGrad
a H caesp
a H caesp/rept
a H caesp
a G rhiz
a Hros
aH ros
aHsem
aHros
T rept
a H caesp/scap
aHsem
a G rad/H scap
T sem
H caesp/G rad
aHsem
aHsem
a H scap
a H ros/G rad
a H scap/G rad
G rad/scap
a H caesp/scap
aH ros/G rad
a H ros/sem
G rad/H sem
a Hros
a H caesp/scap
a H caesp
a H caesp
a Hscap
a H ros/caesp/G rad
aH scap
aHsem
a H caesp/a f Ch
herb pulv
a G rhiz/a f Ch herb
pulv
a Mi P caesp
VI
Vv
VIII
VII, VIII
Ill
Ill
Vv
VIII
Ill, V, VII
VII
|b ag
VIII
VI, VII
VIII
By.41, 213, 374, 892
Mc.294, BL.2048
BL.2026, Mc.292
BL.2025
BL.2124, M.1095, W.230,
L.955, ons.
Z.1842
Onyon 9
By.287, M.1004, P.s.n.,
S.7598, Z.599, Sm.s.n.
S.7109
BL.2107
By.758
Z.449
M.s.n., Sm.s.n.
By.346, M.864, Mc.328
By.518, L.0128, M.900
BL.2184, H.11, Mc.352,
Sm.s.n.
By.s.n., W.423
H.29
BL.2093, By.258, D.351,
W.286
Beer 25339
BL.2030, Mc.268
B.5647, By.371, M.896
H.31, BL.2181, By.277
By.289, 641, M.846
BL.2137
B.5697, By.247, H.1, P.55
BL.2129, Mc.366
D.433
D.411
BL.2086, 2201, S.4603
By.422, Shresta 5661,
Sm.s.n.
BL.2180, By.434, D.381
By.368, D.350, L.0185,
M.965, P.G.234
Mc.285
BL.2088, Mc.388
M.1099, Mc.335, 373
BL.2158
Peracarpa carnosa (Wall.)
Hook.f. & Thoms.
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 (Lind1.) J.
Moore
Pleurospermum angelicoides
(DC.) C. B. Clarke
P. apiolens C. B. Clarke
*
P. benthamii (DC.) C. B. Clarke
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 *
*
T rept/scap
aG bulb
aG bulb
T rept
aH rept
a G rhiz
a NP caesp
a H caesp
aH scap
aHros
a Ch frut/a NP
caesp
aGrad
a G rhiz/ros
s NP scap
a G bulb
a G rhiz
aH ros
b Mes P scap
a Mi P scap
aG bulb
aG bulb
fac a GE bulb
a Grad
a Grad
a Grad
a Grad
a Grad
a Grad
a Grad
a Grad
T caesp
a H caesp
a H caesp
a H caesp
a G bulb
a G rhiz
a G rhiz
PLANTS SOUTH OF MT EVEREST
I
ng.
251
Z.778
H.60
Mc.396
M.917, W.244
Zit3t
P.s.n., Sch.2036, 2039
P-s.n:
L.2089, S.4630, Z.733,
1829
P.G.311, W.428
By.585, P.R.310, Sch.2190,
2280, Z.776
BL.2130, M.1055,
P3G:247., 2.50), 375,
Sm.s.n.
M.819
M.1036
BL.2051, Mc.280
By.911, M.857, P.s.n.,
ZAs19 51937
W.333
BL.2014
Beer 25369, S.660
M.s.n.
By.1175, P.s.n., Sch.2082
H.65
H.66, L.0171
M.s.n. (Gottingen, Bot.
Garden)
F.8-0;
M.910a, 937, 987, 1016,
Psen:
By.644, L.7, M.921
M.847
By.656
By.663, 831
By.449, M.1085, Sm.s.n.
By.663, 831, M.910b
Sm.s.n.
W.243
M.1103, Sm.s.n.
P.G.306
Z.618
§.7124
BL.2066, By.880, M.941,
Sm.s.n.
252
P. cf. kingianum Colett & Hemsl.
4+
P. verticillatum (L.) All.
Ponerorchis chusua (D. Don) S06
*
Potentilla argyrophylla Wall. ex
Lehm. *
P. argyrophylla Wall. ex Lehm.
var. argyrophylla
P. argyrophylla var.
atrosanguinea (Lodd.) Hook.f.
*
P. caliginosa Sojak
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. griffithii 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. MIEHE
a G rhiz
a G rhiz
a G bulb
aHsem
aHsem
aHsem
aHsem
aHsem
aH rept/f Ch herb
pulv
a G rhiz
a Ch frut
aHsem
a H caesp
aHsem
aHsem
aHsem
af Ch herb pulv
af Ch herb pulv
af Ch herb pulv
aHsem
aHsem
aH sem/G rhiz
aHsem
aHros
aHros
aHros
aHros
aHros
aHros
aHros
aHros
aHros
aHros
SY
IX
VI, IX
HV, Vil
VII, VIII, X
VII, X
av, Vo
VII
IX
VIII, IX
LO.
IX
P.s.n.
By.878, M.926, Mc.303,
Z..669
By.226
D.403, S.4774
P.G.255, Z.586
f.M.
Sw.258
D.363, Mc.383, P.35
By.196, D.329, M.874,
P.33, 34, Sch.2035,
W.439, Z.468, 613
D.425, W.438, Sm.s.n.
BL .2159; 2533, 7.21,
D.329, M.852, 1079,
P.G.294, G.724, S.4752,
Sch.2033, 2034, Z.613
M.821, 878, Z.1659
P.G.338
M.888
By.942, D.348
BL.2053, M.974, W.418
By.1097, Mc.251
D.349, M.1038, Z.619,
665a
M.967, 1073
BL.2108, Z.580a
D.364, Z.656
By.233, $.4689, 4787
By.45, 208, M.1025, Z.500,
628, 685, 1610a
Sch.2015
By.417, S.4817, Z.269, 340,
512, 563, Sm.s.n.
BL.2054, Mc.378
Beer 25354, Mc.281
Sch.2012
Beer 25408
BL.2123A, P.G.283,
S.6596, W.261, Z.1508,
Sm.s.n.
M.1059, 1109, Z.1505,
Sm.s.n.
By.129, P.s.n., Sch.2198,
2292, Z.370, 402, 505
§.4530, Z.815c
P. dickieana Watt
P. gambeliana Watt
P. geraniifolia Hook.f. *
P. glabra Klatt
P. glomerata Pax. *
P. gracilipes Craib
P. hookeri Watt
P. listeri King ex Hook.f.
P. macrophylla D. Don *
P. macrophylla var.
moorcroftiana (Wall. ex Klatt)
W. W. Sm. & Fletcher
P. megalocarpa Hara
P. muscoides Hook.f. ex Watt
P. obliqua W. W. Sm. *
P. primulina (Spreng.) Hara *
P. pulchra Watt
P. reticulata Wall.
P. rotundifolia Wall. *
P. scapigera (Hook.f.) Craib
P. sikkimensis Hook.f. *
P. stuartii Wall. *
P. tanneri subsp. nepalensis (W.
W. Sm.) A. J. Richards
P. tenuiloba (Watt) Pax
P. uniflora Klatt
P. walshii Craib *
P. wollastonii Balf.f. *
Prunella vulgaris L.
Prunus rufa Hook.f. *
Pseudostellaria heterantha
(Maxim.) Pax
P. heterantha var. himalaica Ohwi
Pueraria peduncularis (Benth. )
Grah.
Pyrola sikkimensis Kiisa
Quercus glauca Thunb. var.
glauca *
Q. lanata Sm. *
Q. semecarpifolia Sm.
Rabdosia lophanthoides var.
gerardiana (Benth.) Hara
PLANTS SOUTH OF MT EVEREST
aHros
aHros
aHros
a Hros
a Hros
a Hros
aH ros
aHros
aHros
aHros
aHros
af Ch herb pulv
aHros
aHros
a Hros
aHros
aHros
aHros
a Hros
aHros
aHros
T ros
aHros
aHros
aHros
T scap
a Mi P scap
a G bulb
aG bulb
a/e d PL suff
aHsem
s Mes P scap
s Mes P scap
s Mes P scap
aH scap
V, Vil
Ill, V
I, Ill
Vv
TV.
Vil
Il
VIII
VIII, 1X
VI, IX
Ill, 1V, VIII
253
Beer 25352
By.390, BL.2037, Z.815
BL.2173, Mc.390, Z.726,
1747
Z.814
By.352, 649, dH.2983A,
H.57, M.894, 991, Z.816,
Sm.s.n.
S.4526
Beer 25374
S.6975, Z.779
BL.2097, W.278
Z.1463
Beer 25373, BL.2192,
dH.2986, M.984,
P..G.284, 2.1565,
Sm.s.n.
Mc.389
Beer 25353, Mc.382
By.442, BL.2094, D.383,
432, M.972, W.259,
2.5.1: S022, 001-50:
BL.2039, $.7090
Mc.406
By.436, D.361, M.909,
Mc.278, L.0187, Z.506,
636, 1700, Sm.s.n.
Z.800
By.547, 592, 605, 636, 649,
1500, D.369, Sm.s.n.
Mc.386
Z.799
BL.2191
Beer 25360, S.1649
P18:0., Ze282,00l,02e,
Sm.s.n.
BL.2101, 2136, By.274,
D.402, H.18, M.842,
P.R.312, 2.1626, 1807,
Sm.s.n.
By.585, W.414
Sch.2256
By.828
D.344
dH.2911A
Z.483
D.320
S.4539
f.M.
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
R. anthopogon var. hypenanthum
(Balf.f.) Hara
R. arboreum Smith *
R. arboreum var. cinnamomeum
(Wall. ex G. Don) Lindl.
R. barbatum Wall. ex G. Don *
a Hros
a Hros
aH ros
aH ros
aH ros
aHros
a Hros
aHsem
G. MIEHE
a H ros, fac hyd
aH ros
aH ros
aH ros
aH ros
aHros
aH sem/G rad
a Ch 1 succ
a Ch1 succ/G rad
a Ch1 succ
a Ch1 succ/G
rhiz/caesp
a Ch | succ/caesp
a Ch | succ/a f Ch
herb pulv
a Ch] succ
a Ch 1 succ
e Ch! succ
a Ch 1 succ
a Ch1succ
a Ch 1 succ
a Ch1succ
s NP caesp/s Ch frut
pulv
s NP caesp/s Ch frut
pulv
s Mes P scap
s Mes P scap
s Mi P caesp
Ill
VII, VIII
VIII, 1X
I, i
VII, Vill
VIII
VIII
Vv
VIII
VIII
Ill
VIII
IX
IV
IV
VII, VIII
X
Ill, 1V
IV, VIII
VII
Vil
I-III
I-III
I, Il
S.7091
D.338, H.38, Sm.s.n.
W.202
Z.745
W.458, Z.582, Sm.s.n.
Sm.s.n.
BL.2080, 2189, Sm.s.n.
P.R.284
BL.2146, W.197
By.230, BL.2076, M.1113,
1115, P.s.n.
$.7092
Sm.s.n.
BL.2106, D.406, S.7110,
S.7134
By.780
Beer 25514, BL.2154,
By.180, M.1017, 1044,
P.G.233, Sm.s.n.
By.779, M.1063, W.168,
Sm.s.n.
BL.2077
L.0211, Mc.349, S.4615
BL.2084, 2085, 2095,
By.49, M.1105, Mc.339,
Z.1514
By.408, D.378, Sch.2180,
Z.701, Sm.s.n.
M.863
H.13, 68, M.841
D.396, 397
M.1062, 1096
By.702, 793, 794, M.838,
Mc.270, P.103,
Z.1628
H.27, 58, M.923, P.R.301,
s.n.
M.951
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. campylocarpum 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. lindleyiT. Moore *
R. nivale Hook.f. *
R. setosum D. Don *
R. triflorum Hook.f. *
R. wightii Hook.f. *
Ribes glaciale Wall.
R. griffithii Hook.f. & Thoms. *
R. himalense Royle ex Decne.
R. luridum Hook.f. & Thoms.
R. orientale Desf. *
R. takare D. Don *
R. takare forma desmocarpum
Hara
R. vilmorinii Janczew.
Rodgersia nepalensis T. A. Cope
ex Cullen *
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
s Mi P caesp I, Ill
s Mi P caesp III
s NP caesp/sChfrut I, II
s MiP caesp I
s NP caesp/fac E I
s NP caesp/s Mi P Il
caesp
s Mi P caesp Ill
s/a NP caesp Ill, V
s/a NP caesp V, VII
s NP caesp/fac E I
s Ch frut/s Ch frut Vit be
pulv
sNPcaesp/sChfrut VII
pulv
sNPcaesp/sChfrut I, I
s Mi P caesp Ill
a NP caesp Div
a NP caesp Il
a NP caesp I
a NP caesp Vil
a NP caesp I
a NP caesp III
a NP caesp Il
a NP caesp Vv
e Grad II
a NP caesp III, V
a NP caesp Ill, V
a Grad I-III, V
a NP caesp ?
a Ch frut rept Vv
a NP caesp/a PL Vv
frut
a Ch frut rept I, I
ad PL frut I,IV
a Grad i, Vx
ad PL frut I
as
Mc.47, S.4527, Sch.2252
By.100a, s.n., M.856, 931,
Mc.49, Sch.2188, $.2252,
4602, Z.373, 412, 499
Mc.57, $.4613, Z.793
By.186, BL.2031, Mc.277,
S.4532
Z.463
M.930, Mc.288, S.4524
By.685, 686, 968, Mc.48,
$.4521, Sch.2343,
W.411, 2.645
By.184, 296, 385, 659,
1008, 1109, D.347,
M.853, 860, Sch.2281,
Z.673, 1669, 1782,
1783B
Sm.s.n.
Mc.39, $.4531
By.915, D.430, M.996,
Sch.2269, Z.523A, 538,
1484, 1811, Sm.s.n.
By.688, M.950, Sch.2048,
P.s.n., W.442, Z.504,
590, 1810, Sm.s.n.
Mc.40, Sch.2048, S.4520,
Z.431
Z.1784A
M.920
S.4534
Z.414, 462
D.355
Z.616
L.0152, $.4619
By.5, dH.2976, L.0168,
P.37
By.17, dH.2938, M.855,
P.38, 39, R.278, 305
f.M., By.18, 327, 372
By.579
Z.692
dH.2990, M.904, P.40
dH.2931
dH.2931
By.215239),.0 8-0;
Z.1720, Sm.s.n.
BL.2018, Z.481, 489
256
Salix calyculata Hook.f. ex
Anderss. *
. daltoniana Anderss.
. disperma Roxb. ex D. Don
. eriostachya Wall. ex Anderss.
. hylematica Schneid.
. ef. insignis Anderss. +
. lindleyana Wall. ex Anderss.
. longifora Anderss.
. luctuosa H. Lév.
S. serpyllum Anderss.
S. sikkimensis Anderss. *
S. tetrasperma Roxb.
Salix sp.
Salix sp.
Salvia campanulata Wall. ex
Benth. *
S. castanea Diels
AN AN AnNN i)
S. hians Royle ex Benth. *
Sanguisorba diandra (Hook.f.)
Nordborg
Sanicula elata Buch.-Ham. ex D.
Don
Sarocococca hookeriana Baill.
Satyrium ciliatum Lindl.
S. nepalense D. Don *
Saussurea auriculata (DC.) Sch.
Bip. *
S. candolleana (DC.) Sch. Bip.
S. deltoidea (DC.) Sch. Bip. *
S. gnaphalodes (Royle ex DC.)
Sch. Bip. *
S. gossypiphora D. Don *
. graminifolia Wall. ex DC. *
. hieracioides Hook.f.
. leontodontoides (DC.) Sch.
Bip.
. leontodontoides var. filicifolia
(Hook.f.) Hand.-Mazz.
. nepalensis Spreng. *
. simpsoniana (Field. & Gardn.)
Lipschitz *
. tridactyla Sch. Bip. ex Hook.f.
. uniflora Wall. ex Sch. Bip.
. wernerioides Sch. Bip. ex
Hook.f.
S. yakla C. B. Clarke
Saussurea sp.
Saxifraga andersonii Engl. *
HAN AN ~ ~H H i)
G. MIEHE
a Ch frut rept
a Ch frut/a NP
caesp
a Mi P scap
a Ch frut rept
a Ch frut rept
a NP caesp
a Ch frut rept/f pulv
a Mi P scap
a NP caesp
a Ch frut rept
a NP caesp
a NP caesp
a NP caesp
a Mi P scap
aHsem
aH sem
aH sem
aH scap
aHsem
s NP caesp
a G bulb
a G bulb
aHsem
a H scap
a H scap
a G rad/H ros
a H scap/G rad
a H scap/g Ch herb
pulv/G rad
a G rad/H ros
aH ros/G rad/f Ch
herb pulv
aHros
a H ros
a G rad/f Ch herb
pulv/H scap
a H scap/G rad
a H scap/G rad
af Ch herb pulv
H ros/G rad
a Grad/ros
e g Ch herb pulv
VII
Ill, VII
I
VII
VII
IX
Ill, Vil
VI, IX, X
IX, X
B.5678, BL.2134, By.550,
D.387, 388, M.924, 952,
Sm.s.n.
By.6, 446, D.457, M.1003,
W.343, Z.376
Z.471
By.189, 531, W.311
By.543, D.354, 437,
W.431, Z.1518
P.R.344
D.436, M.962, 990, 1005,
Sm.s.n.
Z.387
Z.388, 491
Z.1518
BL.2118, W.419
Z.479
dH.2987
M.922
By.204, S.7141, Z.371A,
374, 407, 654, 1813
By.973, P.s.n., Sch.2055,
2057
BL.2151, By.194, D.447,
M.825
M.871, P.42
D.324
f.M., Sch.2083
By.646, H.64
dH.2881, H.69, P.26
By.622
P.R.352
Pesin:
P.G.236, 256, 296
By.145, M.1000, 1046,
1084, P.G.218, 279,
W.215, Z.1452, Sm.s.n.
M.1061, P.s.n., Z.1450,
Sm.s.n.
P.R.335, W.212, Z.1476,
Sm.s.n.
By.558, 1135, M.953, 995,
1081, W.226, Sm.s.n.
P.s.n. (f.Enum)
Pan:
M.1046A, S.6583, W.214,
Z.1516, Sm.s.n.
M.1047, Z.576, Sm.s.n.
Beer 25390
M.1049, P.G.235, W.217,
Sm.s.n.
P.R.303, W.385
M.919
D.419, 443, Z.566, Sm.s.n.
S. aristulata Hook.f. & Thoms.
S. brachypoda D. Don *
S. brunonis Wall. ex Seringe *
S. cordigera Hook.f. & Thoms.
S. diversifolia Wall. ex Seringe
S. engleriana H. Smith *
S. filicaulis Wall. ex Seringe
S. gageana W. W. Smith
S. georgei Anthony
S. granulifera H. Smith
S. hirculoides Decne.
S. hispidula D. Don
S. hookeri Engl. & Irmsch.
S. jacquemontiana Decne. *
S. lowndesii H. Smith
S. lychnitis Hook.f. & Thoms. *
S. cf. melanocentra Franch. *
S. moorcroftiana (Seringe) Wall.
ex Sternb.
S. mucronulata Royle *
S. mucronulata subsp.
sikkimensis (Hulten) Hara
S. nutans Hook.f. & Thoms.
S. pallida Wall. ex Seringe
S. palpebrata Hook.f. & Thoms.
S. parnassifolia D. Don *
S. perpusilla Hook.f. & Thoms.
S. pilifera Hook.f. & Thoms.
S. pseudo-pallida Engl. & Irmsch.
*
S. punctulata Engl.
S. roylei H. Sm. *
S. saginoides Hook.f. & Thoms. *
S. serrula H. Smith
S. sibirica L. *
S. sikkimensis Engl.
PLANTS SOUTH OF MT EVEREST
g Hherb pulv
e H scap
e Hros
H sem
aHsem
e Hros
e H caesp/scap/f Ch
herb pulv
e Hros
e g Chherb pulv
a Hros
a H ros/f Ch herb
pulv
aH caesp
a H ros
e H ros/g Ch herb
pulv
e H ros/rept/g Ch
herb pulv/G rhiz
af Ch herb pulv
a Hros
a g Ch herb pulv/H
TOs
e Hros
e H ros/rept
aH ros/Ch herb
pulv
aHsem
aHsem
aHsem
e Hros/f Ch herb
pulv
e H ros/rept
a Hros
e Hros
e f Ch herb pulv
e g Chherb pulv
e H rept/caesp
a H ros/sem
a H scap
IV, VI
IV
VI, IX
IV, IX, X
Vv
X
IV, VII
IV, VII, [X
X
Ill, V
IX, X
IV, VI, Ix
IX, X
VIII-x
21
By.673, H.27, 52, M.843,
907, P.G.214, W.211,
Z.1479, 1526, 1615B,
Sm.s.n.
Beer 25412, By.489, 666,
H.7, M.843, Sm.s.n.,
P.s.n., W.287, Z.1876
By.487, M.1018, P.R.350,
W.338, Sm.s.n.
Beer 10003
P.48, Z.1292, 1805
Mc.332, S.7106, Sm.s.n.
By.604, M.865, Z.s.n.
Beer 9581, M.1020
Sw.500
dH.2975, M.898, P.R.340,
W.284, Z.1506, Sm.s.n.
By.542, P.G.s.n., Sm.s.n.
M.908, 1020, P.G.331,
W.260
P.47, S.628, W.374
M.1086
Sw.421
M.1089, Mc.329, Sm.s.n.
By.450, Sm.s.n.
M.1026, Z.1549
By.443, D.417, M.1013,
Z.1615a, Sm.s.n.
BL.2115, Mc.356
H.30, W.207, 287, Z.1526,
1623, Sm.s.n.
BL 2156, By.515,,D.352,
440, L.2202, Mc.297,
S.4780
Beer 9554, D.461
By.391, 781, M.810,
Z.1805
M.1064, 1106, Mc.340,
P°G:237, W232
BL.2143, Sm.s.n.
BL.2095, D.398, 438,
M.1012, Mc.333,
Sm.s.n.
BL.2145, M.1078,
P.G.274, W.180, Sm.s.n.
P.s.n.
BL.2141, By.778, D.413,
H.42, M.1040a, Mc.331,
P.G.238, W.161, 223,
277, Sm.s.n.
P.50
B.5777, By.512
Beer 25512, S.1644
258
S. stella-aurea Hook.f. & Thoms.
S. stenophylla subsp. hoffmeisteri
(Klotzsch) Hara *
S. strigosa Wall. ex Seringe *
Schisandra grandiflora (Wall.)
Hook.f. & Thoms. *
Scopolia stramonifolia (Wall.)
Shrestha *
Scrophularia pauciflora Benth.
Sedum chauveaudii R.-Hamet
S. gagei R.-Hamet
S. oreades (Decne.) R.-Hamet *
S. triactina Berger
S. trullipetalum Hook.f. &
Thoms.
Selinum candollii DC.
S. cortioides C. Norman
S. 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. diversifolius Wall. ex DC.
S. graciliflorus DC. *
S. lancifera J. R. Drummond
S. rufinervis DC.
S. scandens Buch.-Ham. ex D.
Don *
S. triligulatus Buch.-Ham. ex D.
Don
S. wallichii DC.
Sibbaldia cuneata Hornem. ex
Kuntze *
S. micropetala (D. Don)
Hand.-Mazz.
S. perpusilloides (W. W. Smith)
Hand.-Mazz.
S. purpurea Royle *
Silene aff. gonosperma (Ruprt.)
Bocquet
S. gonosperma subsp.
himalayensis (Rohrb.) Bocquet
var. himalayensis
S. nigrescens (Edgew.) Majumdar
S. setisperma Majumdar
S. thomsonii Majumdar
Sinocarum sp.
Skimmia laureola (DC.) Sieb. &
Zucc. ex Walp. *
Smilacina oleracea (Baker)
Hook.f. *
S. purpurea Wall. *
Smilax menispermoidea DC.
Sorbus cuspidata (Spach)
Hedlund *
G. MIEHE
e H ros/rept/f Ch
herb pulv
e H ros/rept
aH sem/rept
ad PL frut
. aH scap
a G rhiz/scap
a Ch 1 succ
a H scap
a Ch | succ
a Ch | succ/caesp
a Ch] succ
a Grad
a Grad
a Grad
aHsem
a Hros
a Ch suff
a H sem/scap
aH scap
a G rad/scap
aHsem
aHsem
a Ch suff/L
a Ch suff
aHsem
ag Chherb pulv/G
rhiz
aHsem
a G rhiz/f Ch herb
ulv
a G rhiz/f Ch herb
pulv
a H caesp
a H caesp
a G rhiz/H sem
a H scap
aH sem
a Grad
s NP caesp
a G rhiz
a G rhiz
sd PL frut
a Mes P scap
X
IV
BL.2145, M.1100, Sm.s.n.
M.1118
By.660, M.906, P.51,
Z.1660
B.5679
Z..1679
H.22, M.1020, Z.1609
By. 620
Mc.270
M.862, 960, W.273, 301
By.13, 245, 655
M.1001a
M.848, 883a
Z.1634, 1770
M.1052
Z.1740A, 1845
By.197, 399
M.827, P.s.n.
Z.1760
M.829
P.s.n.
dH.2909, Sch.2363, W.383,
Z.1740
dH.2901, $.4487
Z.1662
Z.658, 695
BL.2171, D.346, Z.789
Mc.379
M.1039, 1077, Mc.377,
Z.568, Sm.s.n.
M.1027
BL.2187, By.432, Sm.s.n.
M.1006, Sm.s.n.
Sm.s.n.
M.968, P.s.n.
M.s.n.
S.2064, 4477
Mc.267, S.4627
Mc.291
Mc.55, 423, 1783
M.824
S. foliolosa (Wall.) Spach
S. hedlundii Schneid.
S. microphylla Wenzig *
S. ursina (Wenzig) Decne.
Soroseris hookeriana (C. B.
Clarke) Stebb. *
S. hookeriana (C. B. Clarke)
Stebb. subsp. hookeriana
S. pumila Stebb.
Spathoglottis ixioides (D. Don)
Lindl. *
Spiraea arcuata Hook.f. *
S. bella Sims
S. canescens D. Don *
Spiranthes sinensis (Pers.) Ames
*
Stellaria congestiflora Hara
S. decumbens Edgew. *
S. decumbens var. polyantha
Edgew. & Hook.f.
S. decumbens var. pulvinata
Edgew. & Hook.f.
S. graminea L.
S. lanata Hook.f. ex Edgew. &
Hook.f.
S. sikkimensis Hook.f. ex Edgew.
& Hook.f.
Streptopus simplex D. Don *
Sunipia bicolor Lindl.
Swertia cuneata D. Don
S. dilatata C. B. Clarke
S. hookeri C. B. Clarke *
S. kingii Hook.f.
S. multicaulis D. Don
Sympagis petiolaris (Nees)
Bremek.
Symplocos paniculata (Thunb.)
Mig. *
S. sumuntia Buch.-Ham. ex D.
Don
Tanacetum atkinsonii (C. B.
Clarke) Kitam. *
T. gossypinum Hook.f. & Thoms.
ex C. B. Clarke *
Taraxacum mitalii van Soest
T. monochlamydeum
Hand.-Mazz.
T. pseudostenoceras van Soest
Taxus baccata subsp. wallichiana
(Zucc.) Pilger
PLANTS SOUTH OF MT EVEREST
a Mi P scap Ill
a Mi P scap I
a Mi P scap Il
a Mi P scap III
a Hros X
aH ros Xx
a H ros Xx
obl. a E Ch succ IV
a NP caesp III, V
a NP caesp II
a NP caesp II
a Grad Vv
aH rept VIII, X
eHrept/efChherb X
pulv
aH rept xX
af Ch herb pulv/a VIII, X
H rept
aH rept IV, VII
aH rept ?
T rept 4
a Grad Ill
obl. e E Ch succ I
aH sem VII, VIII
aH scap Vv
aH sem/G rad VII
a H sem/G rad Vv
a G rad/H caesp VII
II
s/aMiPscap/caesp_ I
s MiP scap I
aHsem V, VI
a H ros/scap IX, X
a G rad/H ros V-VII
a G rad/H ros Vv
a G rad/H ros V, VI
b N/Mi P scap I
259
M.893
P:8.0:
By.7, 839, 1155, D.451,
S.6580, 7143, 7147
Mc.276
M.998, W.254
P.G.247, Sms: 0.
By.382, P.G.217, Sm.s.n.
By.366, M.806, L.016
By.439, M.933, P.s.n.,
Sm.s.n.
S.4628
S.7122
H.62-64, M.835
W.246
M.1022, Sm.s.n.
BL.2028, By.387, M.971
M.s.n.
M.897
W.373
W.380
By.138, BL.2017, L.0158,
vi 88 f
Sch.2312
M.949, 1057, P.160, R.328
M.918
S.7087
L.0419, W.290
By.504, 524, D.339, H.44,
P.G.209, G.285, W.234,
Sm.s.n.
Z.1743
S.4623
S.4483
P.s.n.
By.1176, H.39, Sm.s.n.,
M.1031, 1070, P.G.211,
G.295, Sch.2315, W.179,
Z.1509
Z.270
Z;,1 163
Z.1603
f.M., Sch.2056, 2939,
S.4496
260
Thalictrum alpinum L. *
T. cultratum Wall. *
T. elegans Wall. ex Royle
T. reniforme Wall. *
T. virgatum Hook.f. & Thoms. *
Thermopsis barbata Royle *
Thesium himalense Royle ex
Edgew.
Tiarella polyphylla D. Don
Tofieldia himalaica Baker
Trachydium aff. roylei Lindl. +
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. &
Thoms. 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
DG.
V. grandiflorum Wall. ex DC. *
G. MIEHE
aHros
aHsem
aHsem
aHsem
aHsem
aGrad
aGrad
aHsem
a G rhiz
aGrad
a H caesp
a H scap
a H caesp
a H scap
a Grad
a H caesp
ael HL
a H caesp
a H caesp
a H ros
b Mes P scap
a G rhiz
a hyd H rept
s E Ch frut
s NP caesp
s fac E Ch frut rept
fac E/s NP caesp/Ch
frut
aH scap
aHsem
a H scap
aHsem
aH scap
aH scap
aH rept/G rhiz
a H scap
a H scap
a Mi P caesp
a NP caesp
D.394, 400, BL.2111,
Z.644, Sm.s.n.
$.7140, Sm.s.n.°
P.s.n., W.405
By.176, 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
P.53, Z.448, 768
M.964
By.561
Beer 10016, S.653
Mc.357, S.7088
BL.2204, S.652
L.0654, P.s.n., Z.679
H.12, W.364
M.808, 817c
Sch.2068
P.G.303
By.1071, M.1021a, P.R.349
BL.2080
f.M., Sch.2340, 2360
By.225
f.M.
P.267
BL.2021
Sch.2163
S.4605, Z.266, 382
P.B.75, S.4475
By.320, D.358, Sch.2077,
Z.1672
Z.263
ZaAisil
By.182, Sch.2061, P.56, 57
P.260
W.191
BL.2176, M.1079, Mc.375,
Sw.88-—99
V. nervosum D. Don *
Vincetoxicum hirundinaria subsp.
glaucum (Wall. ex Wight) Hara
*
Viola biflora L. *
V. pilosa Blume
V. thomsonii Oudemans
Waldheimia glabra (Decne.)
Regel *
Youngia japonica (L.) DC.
Y. racemifera (Hook.f.) Babcock
& Stebb.
Ypsilandra yunnanensis W. W.
Smith & Jeffrey
Zanthoxylum nepalense Babu *
Z. oxyphyllum Edgew. *
PLANTS SOUTH OF MT EVEREST
a Mi P caesp Il
a G rad/scap Vv
a Hros II, III, V
aH ros Ti
aH ros I
a H ros/G rhiz VIII, X
aH sem Sea
a Hsem/a G rad Ill
a Grad V, VII
a/s NP caesp II
ad PL frut/NP II
caesp
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 Fée
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
e rhiz III, V
a caesp I
a rhiz II
a rhiz I
a rhiz I
e caesp I, Il
a caesp Ill
a/e caesp I
a rhiz Ill
a/e caesp I
a caesp I, Il
a rhiz I, Il
a rhiz I
a caesp Vv
a rhiz Ill
a caesp I
a caesp a
a caesp Vv
e rhiz IV
e rhiz IV
e rhiz IV
e rhiz I, Il
261
Mc.35, S.4523, Z.795
Sch.2249, Z.675, 714
By.110, W.366, Z.579, 652,
702, 1774, Sm.s.n.
Onyon 10, $.4479, Z.397
S.4486
By.448, H.36, dH.2999,
M.973, Mc.315, P.R.345,
W.174, 181, Z.1453,
Sm.s.n.
Z.748
M.976, W.319
Beer 25356
Z.158
BL.2015, Z.465, 1695, 1866
Z.1688, 1779
Z.1754
Z.674
dH.2888, Z.256
Z.1871
Z.1896
WsdOd
Z.1019
Z.646
Z.1870
Z.1682
Z.1665
Z.678, 1752
M.885A, 912A, 927
Z.677, 1685, 1794
Z.761
W.354
Z.1823
Lilee
Z.454
Z.1749
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
e caesp
a caesp
a rhiz
a caesp
a caesp
a caesp
a caesp
a caesp
a caesp
a caesp
e caesp
a G rhiz
a rhiz
e/a rhiz
e rhiz
e rhiz
e rhiz
e/a rhiz
a rhiz
b Ch herb rept
b Ch herb rept
a rhiz
a caesp
e rhiz
a rhiz
e caesp
e rhiz
e rhiz
e rhiz
e rhiz
e caesp
e/a rhiz
e rhiz
Ill, V, VII
Vil
Vv
IVS
yy.
VI
VI
II, V1
1 TIE TV.
VII
I
I, Il
1,18
I
Ill
1 BEY
I, WI-—V
PA EV
Vv
M.986, W.256, Z.1562
M.1010A, Z.1618
Z.639, Sm.s.n.
Z.1639
M.885B, 961
M.886
Z.1642, 1684
Z.1681, 1776
Z.273
M.1010B, Z.1542, 1567
Z.385, 488
Z.455, 1832
W.430
LA?
Z.507, 638
Z.311, 396
Z.721
W.387, Z.427, 1890
W.352
Z.703, 1895
dH.2965, M.840, 885c,
925, Z.265, 629, 1650,
1713
Z.754
dH.2882, Z.1658
Z.715
Z.1689B (Candollea 15:
206 [1956])
Z.715B, 1689
W.298, Z.637, 1334, 1624
Z.738
Z.257, 460
2.153
M.912B
PLANTS SOUTH OF MT EVEREST 263
Polystichum acanthophyllum e caesp Ein Z.384
(Franchet) Christ
P. discretum (Don) Diels a caesp I Z.455
P. lachenense (Hook.) Bedd. e caesp VI-X M.1010c, Z.1449, 1595
P. mehrae Fraser-Jenkins & e caesp II N.9
Khullar
P. neolobatum Nakai e caesp I Z.1680
P. nepalense (Spreng.) C. Chr. a caesp I, 11, 1V Z.1742
P. shensoense Christ a caesp III M.975
P. stimulans C. Presl e caesp IV Z.1666, 1737
P. thomsonii (Hook.f.) Bedd. a caesp IV W.356, Z.399, 747, 826b,
1736
Pteridium aquilinum (L.) Kuhn a rhiz II, II, V M.s.n.
Pteris cretica L. e rhiz I Z.1738, 1847
P. dactylina Hook. e rhiz I Z.261 4805-1137
P. subquinata Wall. ex Hope e caesp IV Z.1826
P. wallichiana J. Agardh e caesp 16s Z.1873
Selaginella chrysocaulos (Hook.f. I A We
& Grev.) Spring
Thelypteris levingei (C. B. a rhiz Ill Z.1686
Clarke) Ching
Tricholepidium normale (D. e rhiz IV Z.1750
Don) Ching
Vittaria flexuosa Fée e rhiz IV Z.1748
Woodsia andersonii (Bedd.) a caesp ? W.355
Christ
W. elongata Hook. a caesp IV Z.680, 1751, 1775, 1809
W. lanosa Hook. a caesp Z W.358
Acknowledgements
In the summer and autumn 1982 I had the opportunity to accompany Professor Dr Matthias Kuhle,
Department of Geography, University of Géttingen, 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.
References and other relevant literature
Academia Sinica, Comprehensive Observation Team. 1967. The vegetation of central Tibet. Beijing.
Washington, DC.
Ahrendt, L. W. A. 1956. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie
botanique). 3. — Berberidaceae. Candollea 15: 153-155.
Alston, A. H. G. & Bonner, C. E. B. 1956. Résultats des expéditions scientifiques genevoises au Népal en
1952 et 1954 (partie botanique). 5. — Pteridophyta. Candollea 15: 193-220.
Andrews, C. 1983. Photographs and notes on tourism and deforestation in the Solu Khumbu, Nepal.
Mount. Res. Dev. 3: 182-185.
Baehni, C. 1956. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie
botanique). 2. — Cruciferae. Candollea 15: 149-151.
264 G. MIEHE
—— 1958. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
13. — Juglandaceae, Santalaceae, Thymelaceae et Saxifragaceae. Candollea 16: 215-227.
—— 1959. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
16. — Euphorbiacées et Buxacées. Candollea 17: 69-70.
——., Bonner, C. E. B. & Vautier, S. 1951. Plantes récoltées par le Dr Wyss- -Dunant au cours de
r expédition suisse 4 l’Himalaya en 1949. Candollea 13: 213-236.
Bajracharya, D. 1983. Deforestation in the food/fuel context. Historical perspectives from Nepal. Mount.
Res. Dev. 3: 227-240.
Beer, L. List of seed collections, Kasuwa, Barun, Iswa Khola, autumn 1975. Typescript, BM, London.
——, Lancaster, C. R. & Morris, D. Seed collection Barun, Kasuwa, Iswa Khola, autumn 1971. Typescript,
BM, London.
Bernardi, L. 1963. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie
botanique). 18. - Monochlamydeae, Dialypetalae et Metachlamydeae. Candollea 18: 243-283.
Bjonness, I.-M. 1980a. Animal husbandry and grazing, a conservation and management problem in
* Sagarmatha National Park, Nepal. Norsk geogr. Tidsskr. 34: 59-76.
— 1980b. Ecological conflicts and economic dependency on tourist trekking in Sagarmatha (Mt.
Everest) National Park, Nepal. An alternative approach to park planning. Norsk geogr. Tidsskr. 34:
119-138.
—— 1983. External economic dependency and changing human adjustment to marginal environment in
the high Himalaya, Nepal. Mount. Res. Dev. 3: 263-272.
Boesch, H. 1964. Zwei Jahre Wetterbeobachtungen in Ostnepal (1961-1963). Geographica helv. 3:
170-178.
Bonner, C. E. B. 1959. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie
botanique). 15. — Ericaceae. Candollea 17: 61-67.
Bor, N. L. 1958. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie
botanique). 7. - Gramineae. Candollea 16: 137-142.
Brockmann-Jerosch, H. 1936. Futterlaubbaume und 2 bets apnatinen Ber. d. Schweizer. Bot. Ges. 46:
594-613.
Byers, A. C. & Byers, E. A. 1984. UNN, Nepal Mountain Hazards Mapping Project, Phase II, Khumbu
(Mt. Everest Basin) Nepal. Typescript, Dept. of Geography, University of Colorado, Boulder.
Caine, N., Ives, J. D., Kienholz, H. & Messerli, B. 1982. A buried podsol near Namche Bazar,
Solu-Khumbu, Nepal. Mount. Res. Dev. 2: 405-406.
Cannon, M. J. & Cannon, J. F. M. 1984. A revision of the Morinaceae (Magnoliophyta-Dipsacales). Bull.
Br. Mus. nat. Hist. (Bot.) 12: 1-35.
Chang, D. H. S. 1981. The vegetation zonation of the Tibetan plateau. Mount. Res. Dev. 1: 29-48.
Coburn, B. A. 1983. Managing a Himalayan world heritage site. Nature Resources 19 (3): 229-240.
Dhar, O. N. & Narayanan, J. 1965. A study of precipitation distribution in the neighbourhood of Mt.
Everest. Ind. J. Met. Geophys. 16: 229-240.
Dierl, W. 1978. Grundziige einer 6kologischen Tiergeographie der Schwarmer Ostnepals (Lepidoptera:
Sphingidae). Khumbu-Himal 3: 313-359.
Disselhorst, G. 1968. Beitrage zur Okologie der V6gel Zentral- und Ost-Nepals. Khumbu Himal
2: 1-417.
Dittmann, E. 1970. Statistische Untersuchungen zur Struktur der Niederschlage in Nepal. Khumbu Himal
7: 47-60.
Dobremez, J.-F., Jest, C., Toffin, G., Varnatian, M.-C. &Vigny, F. 1974. Carte écologique du Népal
1:250000. Région Kathmandu- Everest. Paris.
—— & Shakya, P. R. 1977. Carte écologique du Népal. Biratnagar-Kanchenjunga 1:250,000. Paris.
Liste des echantillons collectés au Népal. Mission été 1970. Université Grenoble I, Laboratoire du
Biologie Végétale. Typescript, BM, London.
Dyhrenfurth, G. O. 1959. Die bergsteigerische ErschlieBung der Mt. Everest Gruppe. Jn T. Hagen (Ed.)
Mount Everest. Aufbau, Erforschung und Bevélkerung des Mt. Everest Gebietes: 113-168. Ziirich.
Ellenberg, H. 1954. Steppenheide und Waldweide. Ein vegetationskundlicher Beitrag zur Siedlungs- und
Landschaftsgeschichte. Erdkunde 8: 188-194.
—— 1956. Grundlagen der SAE nS ener ene I. InH. Walter, Einfiihrung in die Phytologie 4: 1-136.
Stuttgart.
— 1966. Leben und Kampf an den Baumgrenzen der Erde. Naturw. Rdsch. Stutt. 19: 133-139.
— 1978. Die Vegetation Mitteleuropas mit den Alpen in 6kologischer Sicht. Stuttgart.
— & Mueller-Dombois, D. 1967.°A key to Raunkiaer plant life forms with revised subdivisions. Ber.
Geobot. Inst. ETH. Stiftung Riibel 37: 56-73.
PLANTS SOUTH OF MT EVEREST 265
Fraser-Jenkins, C. R. 1984. An introduction to fern genera of the Indian subcontinent. Bull. Br. Mus. nat.
Hist. (Bot.) 12: 37-76.
Fiirer-Haimendorf, C. V. 1959. Die Sherpas des Khumbu-Gebietes. Jn T. Hagen (Ed.) Mount Everest,
Aufbau, Erforschung und Bevélkerung des Mt. Everest Gebietes: 169-218. Zurich.
—— 1964. The Sherpas of Nepal. Buddhist highlanders. London.
Funke, F. W. 1969. Religidses Leben der Sherpa. Khumbu Himal 9: 1-369.
Grierson, A. J. C. & Long, D. G. 1983. Flora of Bhutan. Including a record of plants from Sikkim.
Edinburgh.
Gruber, U. F. 1969. Tiergeographische, Skologische und bionomische Untersuchungen an kleinen
Saugetieren in Ost-Nepal. Khumbu Himal 3: 197-312.
Hackel, H., Hackl, K. & Kraus, H. 1970. Tagesgange des Energiehaushaltes der Erdoberflache auf der Alp
Chukhung im Gebiet des Mt. Everest. Khumbu Himal 7: 71-133.
Haffner, W. 1967. Ostnepal-Grundziige des vertikalen Landschaftsaufbaus. Khumbu Himal 1: 389-426.
—— 1979. Nepal Himalaya. Untersuchungen zum vertikalen Landschaftsaufbau Zentral- und Ostnepals.
Erdwissenschaftliche Forschung 12.
Hagen, T. 1969. Report on the geological survey of Nepal. Preliminary reconnaissance. Denkschriften d.
Schweizer. Nat. forsch. Ges. 86 (1).
Hara, H. (Ed.) 1966. The flora of eastern Himalaya. 1. Report. Results of the botanical expedition to eastern
Himalaya organized by the University of Tokyo 1960 and 1963. Tokyo.
— (Ed.) 1968. Photo album of plants of eastern Himalaya. Tokyo.
—— (Ed.) 1971. The flora of eastern Himalaya. 2. Report. Results of the Botanical expedition to eastern
Himalaya organized by the University of Tokyo 1960 and 1963. Tokyo.
——., Chater, A. O. & Williams, L. H. J. 1982. An enumeration of the flowering plants of Nepal 3. London.
——,, Stearn, W. T. & Williams, L. H. J. 1978. An enumeration of the flowering plants of Nepal 1. London.
——, Tuyama, T., Murata, G., Kanai, H. & Togashi, M. 1963. Spring flora of Sikkim Himalaya. Osaka.
—— & Williams, L. H. J. 1979. An enumeration of the flowering plants of Nepal 2. London.
Heuberger, H. 1956. Der Weg zum Tscho Oyu. Kulturgeographische Beobachtungen in Ostnepal. Mitt.
Geogr. Ges. Wien 98: 1-28.
—— 1976. Kulturgeographische Stockwerke und Wanderbewegungen im Hochgebirge am Beispiel des
Mt. Everest-Gebietes. 40. Dt. Geographentag, Tagungsbericht wiss. Abh. 771-774.
Higuchi, K. (Ed.) 1984. Meteorological data in Khumbu and Shorong Himal, east Nepal (revised tables).
J. Jap. Soc. Snow Ice: 132-134.
Hinrichsen, D., Lucas, P. H. C., Coburn, B. & Upreti, B. N. 1983. Saving Sagarmatha. Ambio 12:
203-205.
Holtmeier, F.-K. 1974. Geodkologische Beobachtungen und Studien an der subarktischen und alpinen
Waldgrenze in vergleichender Sicht. Erdwissenschaftliche Forschung 8.
Hornstein, F. v. 1951. Wald und Mensch. Waldgeschichte des Alpenvorlandes Deutschlands, Osterreichs
und der Schweiz. Ravensburg.
Inoue, J. 1976. Climate of Khumbu Himal. Seppyo, J. Jap. Soc. Snow Ice 38: 66-73.
Joshi, D. P. 1982. The climate of Namche Bazar: a bioclimatic analysis. Mount. Res. Dev. 2: 399-403.
Kanai, H., Shakya, P. R. & Shresta, T. B. 1975. Vegetation survey of central Nepal. Jn Ohashi (Ed.), The
flora of eastern Himalaya 2: 415-423. Tokyo.
Kitamura, S. 1967. A list of Compositae collected in Nepal by M. Numata and K. Yoda, Himalayan
expedition of Chiba University, 1963. J. College Arts Sci. Chiba University 5: 85-86.
Kleinert, C. 1983. Siedlung und Umwelt im zentralen Himalya. Jn U. Schweinfurth (Ed.) Geoecological
Research 4. Wiesbaden.
Komarkova, Wald, Jones, Byers & Byers. 1985. Catalog of Nepalese collections, Annapurna, Dhaulagiri,
Everest. Typescript, University of Colorado Herbarium, Boulder.
Kraus, H. 1964. Das Klima von Nepal. Khumbu Himal 1: 301-321.
Kuhle, M. 1978. Obergrenze von Frostbodenerscheinungen. Zeitschr. Geomorph. I, 22: 350-356.
—— 1984a. Auswertungen von Spacelabaufnahmen aus Hochasien. Geogr. Rdsch. 36: 298-301.
—— 1984b. Hanglabilitat durch Rutschungen und Solifluktion im Verhaltnis zum Pflanzenkleid in den
Alpen, den Abruzzen und im Himalaya. Entwicklung und léndlicher raum 18 (3): 3-7.
Larcher, W. 1963. Zur Frage des Zusammenhanges zwischen Austrocknungsresistenz und Frostharte bei
Immergriinen. Protoplasma 57: 569-587.
Lidén, M. 1985. List of Corydalis specimens of the BM. Typescript, Inst. for Systematic Botany, University
of Géteborg, Goteborg.
Limberg, W. 1973. Transhumante Wanderhirten im Nepal-Himalaya. Erdwissenschaftliche Forschungen
5: 182-184.
266 G. MIEHE
— 1982. Untersuchungen tiber Besiedlung, Landbesitz und Feldbau im Solu-Khumbu (Mount Everest-
Gebiet, Ost-Nepal). Khumbu Himal 12: 1-154.
Ludlow, F. & Stearn, W. T. 1975. New Himalayan and Tibetan species of Corydalis (Papaveraceae). Bull.
Br. Mus. nat. Hist. (Bot.) 5: 45-69.
Maie, M. & Shismada, K. 1965. Meteorological data in the Numbur area. Jn M. Numata (Ed.), Ecological
study and mountaineering of Mt. Numbur in eastern Nepal 1963: 124-154. Chiba.
Mehra, P. N. & Bir, S. S. 1964. Pteridophytic flora of Darjeeling and Sikkim Himalayas. Res. Bull. Panjab
Univ. II, 15 (1-2): 69-182.
Meusel, H., Jager, E. & Weinert, E. 1965, 1978. Vergleichende Chorologie der zentraleuropdischen Flora.
Text and maps. 2 vols. Jena.
Miehe, G. 1982. Vegetationsgeographische Untersuchungen im Dhaulagiri- und Annapurna-Himalaya.
Dissertationes Botanicae 66 (1-2). Vaduz.
—— 1984a. Vegetationsgrenzen im extremen und multizonalen Hochgebirge (zentraler Himalaya).
Erdkunde 38: 268-277.
1984b. Waldnutzung im Himalaya, Beispiel Dhauladhar, Himachal Pradesh, Indien. Praxis Geog-
raphie 10: 36-40.
in press. Neue Befunde zur Vegetationshéhenstufung am Mt. Everest. 45. Dt. Geographentag,
Tagungsbericht wiss. Abh. Stuttgart.
Mitsudera, M. & Numata, M. 1967. Meteorology of eastern Himalaya. J. Coll. Arts Sci. Chiba Univ. 5:
75-83.
Miller, F. 1958. Acht Monate Gletscher- und Bodenforschung im Everestgebiet. Berge Welt 12: 199-216.
Murata, G. 1963. A list of plants collected by the Chiba University Rolwaling Himal Expeditions, 1963.
Acta phytotax. geobot. Kyoto 25: 107-118, 26: 77-88, 27: 29-34, 106-113.
— 1965. A list of Labiatae, collected by M. Numata and K. Yoda, Himalayan expedition of Chiba
University 1963. J. Coll. Arts Sci. Chiba Univ. 4: 309.
—— 1967. A List of Sympetalae (exclude Compositae and Labiatae) collected by M. Numata and K.
aot the Chiba University Rolwaling Himal Expedition, 1963. J. Coll. Arts Sci. Chiba Univ.
5: 87-92.
Nishida, M. 1966. Pteridophyta collected by Dr. M. Numata and Dr. K. Yoda, Chiba University Scientific
Expedition to Nepal Himalaya 1963. J. Coll. Arts Sci. Chiba Univ. 4: 571-581.
—— 1968. A supplementary note of Himalayan Pteridophytes. Ann. Rep. Foreign Students Coll. Chiba
Univ. 3: 43-45.
—— 1974. Pteridophyta collected by Mr. P. R. Shakya and Mr. M. Ohsawa, Himalayan expeditions of
Chiba University 1971. J. Coll. Arts Sci. Chiba Univ. B7: 25-33.
Numata, M. 1965a. Grassland vegetation in eastern Nepal. Jn M. Numata (Ed.), Ecological study and
mountaineering of Mt. Numbur in eastern Nepal: 74—94. Chiba.
—— 1965b. Weed flora and communities in eastern Nepal. Jn M. Numata (Ed.), Ecological study and
mountaineering of Mt. Numbur in eastern Nepal: 98-99. Chiba.
—— 1966. Vegetation and conservation in eastern Nepal. J. Coll. Arts Sci. Chiba Univ. 4: 559-569.
—— 1967. Notes on botanical trip in eastern Nepal. J. Coll. Arts Sci. Chiba Univ. 5: 57-74, 243-258.
—— 1983a. Ecological studies in the Nepal Himalayas. Jn M. Numata (Ed.), Biota and ecology of eastern
Nepal: 1-13. Chiba.
— 1983b. Semi-natural pastures and their management. Jn M. Numata (Ed.), Biota and ecology of
eastern Nepal: 455-465. Chiba.
—— 1983c. Structure and dynamics of vegetation in eastern Nepal. Chiba.
Ohashi, H. (Ed.) The flora of eastern Himalaya, 3. Report. Results of the botanical expeditions to eastern
Himalaya in 1972. Tokyo.
——., Ohsawa, M. & Shakya, P. R. 1973. On the occurence of deciduous broad-leaved forests in the cold
temperate zone of the humid Himalayas in eastern Nepal. Jap. J. Ecol. 23: 218-228.
Okuri, J. 1965. List of Cyperaceae and Gramineae collected by M. Numata, Himalayan expedition of
Chiba University. J. Coll. Arts Sci. Chiba Univ. 4: 315-318.
Oppitz, M. 1968. Geschichte und Sozialordnung des Sherpa. Khumbu Himal 8: 1-170.
Pawson, I. G., Stanford, D. D. & Adams, V. A. 1984. Effects of modernization on the Khumbu region of
Nepal: changes in population structure, 1970-1982. Mount. Res. Dev. 4: 73-81.
Poelt, J. Verzeichnis der Aufsammlungen J. Poelt, Nepal 1962. Zusammengestellt von A. Schreiber.
Typescript, Bayerische Staatssammlung Miinchen.
Polunin, O. & Stainton, J. D. A. 1984. Flowers of the Himalaya. Oxford.
Rathjens, C. 1982. Geographie des Hochgebirges. 1. Der Naturraum. Stuttgart.
Rauh, W. 1939. Uber polsterfoérmigen Wuchs. Nova Acta Leopoldina II, 7 (49): 267-508.
PLANTS SOUTH OF MT EVEREST 267
Rolla, S. R. 1963. The Indian Cho Oyo Expedition, 1958: observations of a botanist member. J. Bombay
nat. Hist. Soc. 60: 400-409.
Schilling, A. D. 1975. Everest trek. Typescript, BM, London.
—— 1976. Everest trek. Autumn 1976. Typescript, BM, London.
—— 1977a. Everest trek 1977. Typescript, BM, London.
—— 1977b. Autumn trek 1977. Typescript BM, London.
—— 1978a. Everest trek 1978. Typescript, BM, London.
—— 1978b. The roads to Everest. J. R. hort. Soc. 103: 259-264, 309-314.
—— 1981. East Nepal (Kangchenjunga area) collections. Autumn 1981. Typescript, BM. London.
Schneider, E. 1965. Begleitworte zur Karte Khumbu Himal I und zur Namensgebung. Khumbu Himal 1:
430-446.
Schwarz, R. 1983. Simulationsstudien zur Theorie der oberen Baumgrenze. Erdkunde 37: 1-11.
Schweinfurth, U. 1957. Die horizontale und vertikale Verbreitung der Vegetation im Himalaya. Bonn.
—— 1983. Man’s impact on vegetation and landscape in the Himalayas. Jn W. Holzner, M.J. A. Werger &
I. Ikusima, Man’s impact on vegetation: 297-309. Den Haag.
—— 1984. The Himalaya: complexity of a mountain system manifested by its vegetation. Mount. Res. Dev.
4: 339-344.
Smith, G. F. 1985. Provisional list of plants collected above 4500 m (15000') in Khumbu in July-August
1984. Typescript, Gottingen.
Smith, H. 1958, 1960. Saxifraga in the Himalaya. Bull. Br. Mus. nat. Hist. (Bot.) 2: 83-129, 227-260.
Stainton, J. D. A. 1963. Field notes east Nepal 1963. Typescript, BM, London.
—— 1964. Notes on a journey in east and central Nepal 1964 with brief notes on a journey in east Nepal 1956
and in central Nepal. Typescript, BM, London.
— 1972. Forests of Nepal. London.
Sutton, S. 1978. Plant collectors in Nepal. Jn H. Hara, W. T. Stearn & L. H. J. Williams, An enumeration of
the flowering plants of Nepal 1: 13-21. London.
Swan, L. W. Plants collected in the Barun Khola by L. W. Swan in 1954. Typescript, BM, London.
—— 1961. The ecology of the high Himalayas. Scient. Am. 205 (4): 68-78.
Tamura, M. 1967. Ranunculaceous plants collected by Chiba University Rolwaling Himal expedition in
1963. J. Coll. Arts Sci. Chiba Univ. 5: 141-142.
Teschke, G. C. 1977. Anthropologie der Sherpa. Khumbu Himal 11: 1-183.
Tibet Autonomous Region Scientific and Technical Committee. 1980. An enumeration of the vascular plants
of Xizang (Tibet).
Tilman, H. W. 1938a. Mount Everest. Cambridge.
—— 1938b. The Mount Everest expedition of 1938. Geographical J. 92: 481-498.
—— 1952. Nepal Himalaya. Cambridge.
Troll, C. 1964. Die klimatische und vegetationsgeographische Gliederung des Himalaya-Systems. Khum-
bu Himal 1: 353-388.
Vautier, S. 1956. Résultats des expéditions scientifiques genevoises au Népal en 1952-1954 (partie
botanique). 6. — Polygonaceae. Candollea 15: 221-228.
—— 1959. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
14. Labiatae. Candollea 17: 41-52.
Walter, H. & Walter, E. 1953. Das Gesetz der relativen Standortskonstanz, das Wesen der Pflan-
zengesellschaften. Ber. dt. bot. Ges. 66: 227-235.
—— & Breckle, S. W. 1983. Okologische Grundlagen in globaler Sicht. Stuttgart.
—— —— 1984. Spezielle Okologie der tropischen und subtropischen Zonen. Stuttgart.
— & Lieth, H. 1966. Klimadiagramm-Weltatlas. Jena.
Weibel, R. 1956. Résultats des expéditions scientifiques genevoises au Népal en 1952-1954 (partie
botanique). 4. — Primulaceae. Candollea 15: 157-165.
—— 1958a. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
8. — Crassulaceae. Candollea 16: 143-145.
—— 1958b. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
9.-— Compositae. Candollea 16: 179-194.
—— 1958c. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
10. — Violaceae. Candollea 16: 195-196.
—— 1958d. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
11. — Caprifoliaceae. Candollea 16: 197-199.
— 1960. Résultat des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
17. — Melastomaceae, Myrsinaceae et Campanulaceae. Candollea 17: 81-85.
268 G. MIEHE
Weigel, J. 1969. Systematische Ubersicht tiber die Insektenfresser und Nager Nepals nebst Bemerkungen
zur Tiergeographie. Khumbu Himal 3: 149-196.
Weischet, W. 1965. Der tropisch-konvektive und auBertropisch-advektive Typ der vertikalen Nieder-
schlagsverteilung. Erdkunde 19: 6-14.
Wraber, T. 1972. Determination list (Barun Khola, Dudh Kosi). Typescript, BM, London.
Yoda, K. 1967, 1968. A preliminary survey of the forest vegetation of eastern Nepal. J. Coll Arts Sci. Chiba
Univ. 5: 99-140, 277-302.
Zimmermann, A. 1953. Pflanzen an den obersten Grenzen der Vegetation. Berge Welt 8: 130-136.
—— 1955. Expedition Gaurisankar 1954. List of collected plants. Typescript, BM, London.
—— 1956. Résultats des expéditions scientifiques genevoises au Népal en 1952 et 1954 (partie botanique).
1. —Itinéraires. Candollea 15: 127-147.
Maps:
Khumbu-Himal (Nepal) 1:50000. 2.Ed. 1978 Freytag-Berndt & Artaria: Wien. (= Forschungsunterneh-
men Nepal Himalaya. E. Schneider, F. Ebster).
Shorong/Hinku 1:50000. 2.Ed. 1979. Freytag-Berndt & Artaria: Wien. (= Nepal Kartenwerk der
Arbeitsgemeinschaft fiir Vergleichende Hochgebirgsforschung 5).
Khumbu Area, Nepal. Mountain Hazards Index Map. 1: 50000. 1985. Hallwag: Bern. (The United
Nations University, Highland-Lowland Interactive Systems Project. Field survey by A. Zimmermann, M.
Bichsel, H. Kienholz).
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British Museum (Natural History)
Further genera of the Biddulphiaceae
(diatoms) with interlocking linking
spines
Robert Ross & Patricia A. Sims
Botany series Vol 16 No 4 26 November 1987
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Vol 16 No 4 pp 269-311
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London SW7 5BD Issued 26 November 1987
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
Sa a RP ED ee aR OT aie cape Oe TP LN Ras AMR FELONS MAE Can The ARMAND 269
| WE UA GeO Yer CY et 7 DN A Pe eaters on te alae Aner a Uae Rien Oa EE AA acs One mR 00 NE Sy. od Coe a 269
PER OT SOGCIERE NS BBO TECOLOS 555 oi. 25 oot 50s 0 alien 354344 cahawegcaii aun tee ges teamed ecsaes cans 270
CUS git te a Rie Bre SS a deh CAA AR SAPPY OPE TE ORR yote REED Peete ORT Ente seiner a oF rf
Taxonomic account
IRR 8 5 Ty ae aan a AiO See BP ed eee Oa PR ERERET Cotte ore PASE FORAY Ee ae enor 271
II. Pseudorutilaria (Grove & Sturt ex De Toni & Levi)
RSOVe A StUTCEN Loe 0 OMe ae eae eo an eect Pe es 276
PEE OM ONE ROSSER: Ae SIONS Ooo ice EN ee ee 284
PV Maluingd R Ross Ss Pe Ae SUNS ooo rhwicise ates era a et a 285
Vat TROVE Tes IRAISS Ge Bets SIONS ceo fa te ae ec eae Sa LUA cud ater uns 286
Vir Dextradonaior Ro Ross &.P.. As Sis ecceres hes as ee i es ee 288
TDP SANG SATA OS 0 9 Gk os cited saat fori uidlectene eset yruR es AR Ue UN alan TOR ay te ae meee nets 289
PCM IIR TOCCOA ES isha iia fossa vai ani cee aia e sna alee cboald cate ipa kc dare ewien a eas suse ae 292
Ur go) RA Sear ne ae 8 Ee Pe REIT SR aE TaN oA RE ORS Rh RRS REAR Oe RP at 292
POORER OUMONS OF PALE cree coo cote ee ronope es engin at ose sok SETH Coa reuhon a etter elas da nem Saniee otic 295
ir 1 Ea! ARR kg SO EER CRS Lee vat oir Om MA oe ye aye et err AMAT Rs ery nt mee mete aren nen 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 S. 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 Dextradonator R. 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, Glynggre, Skive, Limfjord, Island of Mors,
Nykjobing, Ejerslev, Island of Fur.
France
Saint Laurent-la Vernéde, Gard
The Miocene deposit from this locality is described by Lefébure (1935). We are not aware of any
more precise information about its geological age.
GENERA OF THE BIDDULPHIACEAE PAB
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.
‘Lamont E 8’
This is the only label on one sample received from Mr Brigger. Neither the California Atademy
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 in 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: pl. 1 fig. 2), Briggera R. Ross & P. A.
Sims (1985: pl. 5 fig. 6), and Trigonium Cleve (Ross & Sims, 1971: pl. 2 fig. 6). On the other hand
there is no thickened ring of silica around these clusters of small areolae such as surrounds the
ocelli of Amphitetras Ehrenberg (Ross & Sims, 1971: pl. 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 ‘antiligulae’. 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. SOLIUM 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 linking 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.the 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
Tempére & 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 Gombos (1982).
1. Solium exsculptum Heiberg, Consp. crit. Diat. Dan.: 52 (1863).
(Pl. 1, Pl. 12 figs 86-88)
Hemiaulus exsculptus (Heiberg) Schiitt in Engler & Prantl, Naturl. Pflanzenfam. 1 (1b): 97 (1896).
Trinacria exsculpta (Heiberg) Hustedt in Rabenhorst, Krypt.-Fl. Deutschl., Osterr. u. d. Schweiz 7 (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 wm, maximum diameter of 5-polar specimens 35—45 um, height at
centre of valve 7-5—16 um; 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 Paes
their tops 18—38 wm. Areolae poroid, 0-3—0-7 ~m 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 wm 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 wm, 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 wm in diameter, surrounded by a narrow hyaline unthickened rim,
their areolae in radial rows, c. 32 in 10 wm. 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 wm long; c. 6 short spines c. 0-5 wm tall on the
distal side of the tips of the elevations. One labiate process on the central portion of the valve 3-5
jm 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 inclined 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).
Glyng¢gre, Jutland, Denmark (BM 30847).
Skive, Jutland, Denmark (BM 14744, 69223; Tempére & 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 Hd1/12; Heiberg, 1863; Kitton, 1870; Cleve & MOller, 1878;
Tempére & Peragallo, 1889b, 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 Hd1/13;
Heiberg, 1863; Kitton, 1870; Tempére & Peragallo, 1889a, 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 mdepth. Deep Sea
Drilling Project site 340 (Dzinoridze et al., 1978).
Chelyabinsk oblast, U.S.S.R. (Proshkina-Lavrenko et al., 1949b).
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,
CB15.368-—375, 4106-4116; Cheneviére, 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 Kara Tau, 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 Jousé* (Figs 4, 87)
Solium exsculptum [‘exculptum’| var. pentagonalis Kitton ex Walker & Chase, Notes on some new and rare
diatoms [I]: 5, pl. 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-
212).
Although both H. L. Smith (1872) and Van Heurck (1896) included Solium in Hemiaulus,
neither of them published the combination Hemiaulus exsculptus; this was first done by Schitt
* 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 vaiidly 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 215
(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. Oligocene’ 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 Jousé (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: pl. 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 sibling 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 light 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 Glyng¢gre, Jutland, Denmark, that have the
appearance of being initial valves. These have sides of 91 wm and 94 wm. 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.
(1949b) 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., isa
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 peculiar distribution, in low latitudes in the Paleocene
and in high northern latitudes from the lower Eocene to the Oligocene, seems more likely to bea
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. PPEUDORUTILARIA (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni, Syl.
Alg. 2: 854 (1894).
Subgen. Pseudo-rutilaria Grove & Sturt in J. Quekett microsc. Club I, 2: 324 (1886), nom invalid (Arts
9 a ae le
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.
Sit),
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 /nternational 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 aterm. . . denoting its rank’ (Art. 21.1) and that ‘a combination (autonyms excepted) is not
validly published 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 ME
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), Tempére & Peragallo (1890), and Schmidt (1893) used the name Pseudorutilaria monile
earlier than this but did not give the generic name valid 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 of 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 solid 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 Hajés (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
a ae vO ER rsp ge) ia toi cs vg Sem cana idceiealgtc uxt4 tes Weta s eaeeeron. Wuevaded woe cate: 6. hendeyi(p. 283)
1b. Valves bipolar: ;
2a. Central portion pentagonal and with acrenate margin....................0656 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
SAR ER Ne eee ats ye eI eae ARSED UR SEPT y BRE EAS SRE TENA P 1. monile(p. 278)
4b. Margins of the projections undulate, undulations more widely spaced than the
EE ANSVOISE COSUAC: stn. fone vs aes oy avs raes seas cease hoeee ees ee rere 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
BN en, Cee oe Reno Oe a ERT ee eC NTT TURNS Pe ERATE 2. clavata(p. 279)
5b. Gibbosity of the margins of the projections about midway between the central
PORTON ANG TNE APOK aos csc ions os scan gcatie oes vic ches resem ained tascen 3. nodosa (p. 280)
1. Pseudorutilaria monile (Grove & Sturt ex De Toni & Levi) Grove & Sturt ex De Toni, Syll.
Alg. 2: 854 (1894).
(Pl. 2, Pl. 12 fig. 89)
‘Pseudo-rutilaria’ monile Grove & Sturt in J. Quekett microsc. Club II, 2: 324, pl. 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 wm long, 17-24 wm 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 uw 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 wm. Linking spines 3-4 on each elevation, extending rather more than
half-way around the summits, c. 3 wm tall. Areolae poroid, with a slightly raised external rim,
diameter c. 0-3 wm, 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. 8in 10 um. 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 Drilling 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 Ht1/1, Ht1/2, Ht1/3
(Jackson’s Paddock); Grove & Sturt, 1886; Schmidt, 1893; Tempére & Peragallo, 1890 (Bain’s Farm,
Totara); Tempére & Peragallo, 1908 (Bain’s Farm); Laporte & Lefébure, 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 Oligocene. South-western Atlantic, Falkland Plateau, 51° 00-28’ S, 46° 58-30’ W.,
2589 m depth. Deep Sea Drilling 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 implied 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 um 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.
(Pl. 3; Pl. 12 figs 90, 91)
Pseudorutilaria monile sensu Hajos in Init. Rep. Deep Sea Drilling Proj. 35: 829, pl. 14 figs 10-12
(1976), non Grove & Sturt.
Valva bipolaris, recta, 65-155 wm longa, 22—30 um 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 wm distantes, alii 6-8 wm
distantes. Altitudo ad vertices elevationum 12-14 um. Spinae ligantes in utraque elevatione 4-5, circum
paullo plus quam dimidium proximalem verticis positae, supra valde expansa, 2-0-—3-5 xm altae. Areolae
poroides, volis occlusae, extrinsecus margine elevato, diametro c. 0-3 wm, in fronte et parte superiori limbi
irregulariter dispersae, 6—8 in 10 wm sed in portione centrali distantiores, in latere proximali elevationum
carentes vel rarissimae. Areolae subocellorum 40-50 in 10 wm. Annulus ex c. 6 processibus occlusis
constans in portione centrali, in valvis curtissimis carens. Spinae marginales acutae, eis valvae continguae
non oppositae, c. 5 in 10 wm. 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 wm alta, seriebus verticalibus areolarum parvarum c. 20 in 10 wm, areolis c. 15 in 10 wm.
Typus: BM 81136, ex stratis eocaenicis de profundis maris Atlantici australis.
Valve bipolar, straight, 65-155 wm long, 22-30 wm 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 xm apart, the others 6-8 um apart. Height to the top of the elevations 12—14 wm. 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 um tall. Areolae poroid,
occluded by volae, with a slightly raised external rim, diameter c. 0-3 wm, scattered over the
valve surface and the upper part of the mantle, 6-8 in 10 wm, 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 wm. 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 wm. 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 um deep, with vertical rows of areolae, rows c. 20 in 10 wm,
areolae c. 15 in 10 wm.
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 (Hajés, 1976).
South-western Pacific, Tasman Sea, 43° 54-60’ S., 154° 16-96’ E., 4750 m depth. Deep Sea Drilling
Project site 283 (Hajos, 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 Hajés’s figures (1976: pl. 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 wm 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-like 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.
(Pl. 4; Pl. 12 fig. 92)
Valva bipolaris, recta curvatave, 145-195 wm longa, 32-34 wm lata; portio centralis circularis, ab
projecturis costis internis transversis separata; projecturae costis transitae, apices paullo clavatos rotunda-
tos versus gradatim angustatae, sed medio gibbosae, marginibus alibi integris. Costae 7-10 wm distantes,
eae medio propriores distantiores. Altitudo ad vertices elevationum c. 15 wm. Spinae ligantes 2—4 in
utraque elevatione, circum dimidium proximalem verticis positae, c. 4 wm altae. Areolae poroides,
extrinsecus margine paullo elevato, diametro c. 0-3 wm, in fronte et parte superiori limbi irregulariter
dispersae, c. 6 in 10 wm. Areolae subocellorum 40-45 in 10 wm. Annulus ex 6-7 processibus occlusis
constans in portione centrali. Spinae marginales acutae, eis valvae continguae non oppositae, 5-7 wm
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 um long, 32-34 wm 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 wm apart, those near the apices the more closely spaced. Height
to the top of the elevations c. 15 wm. Linking spines on each elevation 2—4, extending half-way
around the summit on its proximal side, c. 4 wm tall. Areolae poroid, with a slightly raised
external rim, diameter c. 0-3 ~m, scattered over the valve face and the upper part of the mantle,
c. 6in 10 wm. Areolae of the subocellus 40—45 in 10 wm. Ring of 6-7 occluded processes on the
central portion. Marginal spines pointed, not opposite those of the sibling valve, 5-7 wm 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 CB11.813-814, 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
swelling 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.
(Pl. 5; Pl. 12 fig. 93)
Valva bipolaris, curvata, 150 wm longa, 32 wm 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 um distantibus. Costae c. 8 wm distantes. Altitudo ad vertices elevationum c. 13-5 wm. Spinae
ligantes 2—3 in utraque elevatione, circum dimidium proximale verticis positae, supra valde expansae, c. 4
pm altae. Areolae poroides, extrinsecus margine paullo elevato, diametro c. 0-5 wm, in fronte et parte
superiori limbi irregulariter dispersae, 6-8 in 10 wm, in portione centrali distantiores. Annulus ex 3-4
processibus occlusis constans in portione centrali. Spinae marginales 2—3 in 10 wm, 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 nonnullis carentes.
Typus: BM 81300, ex stratis eocaenicis de profundis maris Atlantici australis.
Valve bipolar, curved, 150 wm long, 32 um broad; central portion circular, separated from the
projections by transverse costae; projections crossed by costae, at their proximal ends about *4
the width of the central portion, tapering to narrow rounded apices, their margins undulate, not
crenulate, the crests of the undulations 15—18 wm apart. Costae c. 8 wm apart. Height to the top
of the elevations c. 13-5 wm. Linking spines 2—3 on each elevation, extending half-way round the
proximal side of the summit, strongly expanded above, c. 4 wm tall. Areolae poroid, with a
slightly raised external rim, diameter c. 0-5 ym, scattered over the valve face and the upper part
of the mantle, 6-8 in 10 wm, 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 sibling valve, 2-3 in 10 wm, 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.
(Pl. 6; Pl. 13 figs 94, 95).
Eunotogramma weissei var. productum [‘producta’] sensu Fenner in Init. Rep. Deep Sea Drilling Proj.
39: 519, pl. 27 figs 3-5 (1977), non Grove & Sturt.
Valva bipolaris, angulo 30°—45° inter projecturas, margine ubique crenulato, 50-155 wm longa, 25-45 wm
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
um distantes. Altitudo ad vertices elevationum 8-5—11 wm. Spinae ligantes in utraque elevatione 1-2,
plerumque supra expansae et implexae, c. 2 wm altae, aliquando spina una decrescens c. 3-5 um alta.
Areolae poroides, volis occlusae, extrinsecus margine elevato, diametro c. 0-3 yw, in fronte et parte
superiori limbi irregulariter dispersae, 6-10 in 10 wm. Areolae subocellorum c. 50 in 10 wm. 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 superiori 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 wm long, 25-45 um 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 wm. Linking spines 1-2 on each elevation, normally
expanded above and interlocking, c. 2 wm tall, sometimes a tapering spine c. 3-5 um tall.
Areolae poroid, occluded by volae, with a raised external rim, diameter c. 0-3 wm, scattered
over the valve surface and the upper part of the mantle, 6-10 in 10 um. Areolae of the subocellus
c. 50 in 10 wm. 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, slightly 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 Atlantic, 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 hyaline area
surrounded either by eight or nine low cuneate spines or by a low flange-like 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 wm 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
pm 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 wm, 60 wm, and 80 um 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.
(Pio 73-PE AS he. 96)
Valva tripolaris, marginibus concavis, crenulatis, 95-185 wm 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 wm distantes. Crenae marginis c. 11 zm distantes, eae in lateribus oppositis projecturae
non semper oppositae. Altitudo ad vertices elevationum c. 10 wm. Spinae ligantes 3—4 in utraque
elevatione, supra expansae et implexae. Areolae poroides, diametro c. 0-3 wm, in fronte irregulariter
dispersae vel in seriebus obliquis indistinctis positae, 6-8 in 10 wm, prope centrum valvae et in parte
superiori limbi distantiores vel carentes. Area irregularis diametro c. 10 ~m ad centrum valvae porellis c.
30 in 10 wm instructa. Annulus ex c. 12 processibus occlusis constans ad centrum valvae, aream porellorum
cingens. Spinae marginales c. 4 wm distantes, irregulariter positae, eae valvae contiguae non oppositae,
supra dendriticales et hic illic implexae. Rimoportula una vel duae approximatae in annulo picecssuua
occlusorum posita.
Typus: BM 81142, ex stratis eocaenicis de profundis maris Atlantici australis.
Valves tripolar, with concave crenulate margins, 95-185 um 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 wm apart. Crenulations of the margin c. 11 wm apart, not always opposite on
the two sides of a projection. Height to the top of the elevations c. 10 wm. Linking spines 3—4 on
each elevation, expanded above and interlocking. Areolae poroid, diameter c. 0-3 um,
scattered or in indistinct oblique rows on the valve surface, 6—8 in 10 wm, sparser to absent near
the centre of the valve and on the upper part of the mantle. An irregular area, diameter c. 10 um,
of porellic. 30 in 10 um 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 wm apart, irregularly spaced, not
opposite those of the sibling 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.
Ill. 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,
centro 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 & 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 earlier 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.
(Pl. 8; Pl. 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 wm longa, 20-25 um 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 wm. 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 wm, in fronte, in elevationibus et in parte superiori limbi ubi ambitus valvae convexus est irregulariter
dispersae, 4—6 in 10 wm; areolae subocellorum 15-20 in 10 wm, diametro c. 0-1 wm. Pori interstriales in
fronte irregulariter dispersi. Spinae superficiales ad 4 wm 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 wm
minimum alto et, pro parte maxima, basi fultura munito constans. Taenia sola visa c. 8 wm alta, ad unum
apicem antiligula valida instructa, seriebus verticalibus areolarum ornata, seriebus areolisque 10-12 in 10
pm.
Typus: BM coll. Adams H941, specim. I, 12, ex depositione miocenica ad ‘St. Laurent-la-Vernéde’, Gard,
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 wm long,
20-25 wm 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 um. 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 wm,
scattered over the valve surface, the elevations, and the upper part of the mantle where the valve
outline is convex, 4—6 in 10 wm; areolae of the subocelli 15-20 in 10 wm, diameter c. 0-1 wm.
Scattered interstrial pores on the valve surface. Superficial spines up to 4 wm 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 wm tall, mostly with a single
buttress at the base. One girdle band seen, about 8 xm deep, with a well developed antiligula at
one apex and with vertical rows of areolae, rows and areolae 10-12 in 10 um.
Miocene. St Laurent-la-Vernéde, Gard. 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 summum utraeque
elevationis in latere suo distali subocello minimo. Limbus verticalis. Areolae poroideés, parvae, in partibus
valvae sparsae carentesve. Spinae ligantes 2—4 in latere proximali 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.
(Pl. 9)
Hemiaulus centralitenuis R. Ross & P. A. Sims in Beih. Nova Hedwigia 54: 182, pl. 2 figs 7-9 (1977).
Valves elliptical, 20-55 um long, 15-25 um broad, 10-12 wm 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 wm in diameter; height to the top of the elevations 15—25 wm. 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
jzm 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 wm, areolae 30-35 in 10 wm; areolae also dense, 25—30 in 10 wm,
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
porellic. 60in 10 wm. 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 um 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,
Sims & 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 linking 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.
(Pl. 10; Pl. 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;
longitudo lateris 115-125 wm. Elevationes sub-triangulares, interdum a limbo depressione angusta
horizontali separatae. Altitudo ad vertices elevationum c. 30 wm. 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 wm altas, c. 5 wm 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 wm, in fronte in seriebus radialibus
dispositae, seriebus 2-5—4 in 10 wm, areolis 2—4 in 10 wm, 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 ~m; in limbo prope marginem series una horizontalis areolarum 2-5-3 in 10 wm
et, prope cristam marginalem, alter series similaris. Pori interstriales in fronte irregulariter 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 hanc interruptus; areolae pseudocellorum in seriebus dispositae, seriebus in vertices elevationum
plus regularibus quam alibi; series areolaeque 8-10 in 10 um; areolae in vertices elevationum diametro
0-2—0-4 wm, minor quam eae alibi diametro 0-5—1-0 wm. Rimoportulae 2-4, prope aream centralem
hyalinam irregulariter dispositae, unaquaeque apertura interna recta, curta, paullo elevata, extrinsecus ex
tubo verticali c. 6 4m alto appendice acuta lanceolata ad apicem instructo constans. Valvocopula parte
exteriori c. 20 wm alta, ad omnes apices antiligula rotundata humili instructa, et areolis irregulariter
dispersis 2-5—4 in 10 wm ornata; pleura parte exteriori c. 10 wm alta et parte interiori hyalina c. 4 wm alta,
pars exterior serie areolarum c. 5 in 10 wm secus marginem proximalem et, huic distalibus, seriebus duabus
paullo irregularibus areolarum c. 4in 10 wm.
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 wm. Elevations sub-triangular, sometimes
separated from the mantle by a narrow horizontal depression. Height to the summit of the
elevations c. 30 wm. Linking spines 15—20 on each elevation, expanded and rather irregularly
shaped above, 2—4 um tall. A low hyaline marginal ridge running from elevation to elevation
and bearing acute marginal spines 10-12 wm tall and c. 5 wm 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 wm; radial rows of areolae on
the valve face, rows 2:5—4 in 10 um, areolae 2—4 in 10 wm, 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 «wm in diameter.
Mantle with a horizontal row of areolae 2:5—3 in 10 wm 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 um;
areolae on the summits of the elevations 0-2—0-4 wm in diameter, smaller than those on the distal
face and the mantle, which are 0-5—1-0 wm in diameter. Labiate processes 2—4, irregularly
distributed a little outside the central hyaline area, with a slightly raised, short, straight internal
opening and, externally, a vertical tube c. 6 wm tall with a lanceolate acute appendage on its tip
c. 2 um long. Valvocopula with a pars exterior c. 20 wm deep, with a slightly rounded antiligula
at each apex and irregularly scattered areolae 2-5—4 in 10 xm; pleura with a pars exterior c. 10
wm deep and a hyaline pars interior c. 4 um deep, the pars exterior with a row of areolae c. 5in 10
um along its proximal edge and two somewhat irregular rows c. 4 in 10 wm 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—Oligocene boundary (Holmes & Brigger,
1979). There is no firm date available for the material from Springfield. We have examined the
holotype of 7. exornatum (BM 3079) and a number of other specimens of the species with the
light 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 wm on the
parts of the valve surface between the centre and the elevations, whereas those of B. simulans
are 3—4 in 10 wm 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.: pl. 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 xm, height to
the centre of the valve 20-30 um. Elevations two, marginal, cylindrical, 5-12 wm in diameter,
height to the top of the elevations 55-75 wm. A hyaline marginal ridge c. 10 wm tall and turned
inwards along its upper edge extending from elevation to elevation. Mantle vertical, hyaline.
Areolae poroid, c. 0-8 ~m in diameter, vela eroded on all specimens seen, in radial rows on the
valve surface, rows and areolae 9-10 in 10 wm, extending up the proximal side of the elevations;
a few small areolae or pores, 0-2—0-4 um 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 zm. Two flattened, acute linking spines on each elevation, partially encircling the
elevation of the sibling valve, those on one ofa 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 wm
tall. No labiate process seen. Cingulum consisting of two entire bands; the valvocopula 60-75
um 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 ~m in diameter and 3-5 wm apart; pleura hyaline, 15—30 wm 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-D1 (BM SEM B26.387-—392, CB11.301—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 um,
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 um 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 anumber 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, Coscinodiscus, 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 asomewhat 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 /sthmia 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 Rattrayella 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: pl. 1 fig. 2) and Biddulphiopsis (see von Stosch & Simonsen, 1984), but
also Hemiaulus (see Ross, Sims & Hasle, 1977: pl. 5 fig. 32, pl. 8 fig. 52), Trigonium (see Ross &
Sims, 1971: pl. 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 ocelli 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 ocelli 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 J. W. Bailey, nom. cons., published in 1851.
However, the family name was published by Kitzing in 1849 and is based on Eupodiscus
Ehrenberg, nom. rej., the type of which is Eupodiscus germanicus (Ehrenberg) Ehrenberg, a
synonym of Aulacodiscus 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 sibling 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 porelli 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 small 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 S. inclinata
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: pl. 21 fig. 5, pl. 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, Dicladiopsis 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
Jousé, 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 valvocopula (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 fiir 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.
References
Bailey, J. W. 1851. Microscopical observations made in South Carolina, Georgia and Florida. Smithson.
Contr. Knowl. 2 (8): 1-48, pl. 1-3.
Benda, L. 1972. The diatoms of the Moler Formation of Denmark (Lower Eocene). A preliminary report.
Beih. nov. Hedwigia 39: 251-266.
Cheneviére, E. 1934. Sur un dépét fossile marin a diatomées situé 4 Kamischev (Russie centrale). Bull.
Soc. fr. Microsc. 3: 103-107, pl. 5-8.
GENERA OF THE BIDDULPHIACEAE 293
Cleve, P. T. & Moller, J. D. 1878. Diatoms. III Part, 9 pp. Uppsala.
De Toni, G. B. 1894. Sylloge Algarum omnium hucusque cognitarum. 2 Bacillarieae. Sect. III.
Cryptorhaphideae. pp. 819-1556. Patavii.
—— & Levi, D. 1887. Algae novae. Notarisia 2: 333-353.
Dzinoridze, R. N., Jousé, A. P., Koroleva-Golikova, G. S., Kozlova, G. E., Nagaeva, G. S., Petrushevs-
kaya, M. G. & Strel’nikova, N. I. 1978. Diatom and Radiolarian Cenozoic Stratigraphy, Norwegian
Basin; D.S.D.P. Leg 38. Initial Rep. Deep Sea Drilling Proj. 38: 289-427.
& Strel’nikova, N. I. 1979. Opisanie Diatomovykh Vodoroslei [Descriptions of diatoms.] In
Istoriya Mikroplanktona Norvezhskogo Morya (po materialam glubokovodnogo bureniya). /ssled.
Fauny Morei 23 (3): 32-70.
Ehrenberg, C. G. 1839. Hr. Ehrenberg las tiber jetzt wirklich noch zahlreich lebende Thier-Arten der
Kreideformation der Erde. Ber. Akad. Wiss. Berlin 1839: 152-157.
—— 1840. Uber noch zahlreich jetzt lebende Thierarten der Kreidebildung. 94 pp., 4 Taf. Berlin & Leipzig.
Reimpr. in Phys. Math. Abh. K. Akad. Wiss. Berlin 1839: 81-174, Taf. I-IV (1841).
—— 1844. Zum Schluss legte Hr. Ehrenberg der Akademie einige verlaufige Resultate seiner Unter-
suchungen der ihm von der Siidpolreise des Capitain Ross, so wie von den Herren Schayer und Darwin
zugekommenen Materialen tber das Verhalten des kleinsten Lebens in den Oceanen und den gréssten
bisher zuganglichen Tiefen des Weltmeers vor. Ber. Akad. Wiss. Berlin 1844: 182-207.
Fenner, J. 1977. Cenozoic diatom biostratigraphy of the equatorial and southern Atlantic Ocean. Jnitial
Rep. deep Sea Drilling Proj. 39: 491-623.
—— 1985. Late Cretaceous to Oligocene planktic diatoms. Jn H. M. Bolli, J. B. Saunders & K.
Perch-Nielsen (Eds.), Plankton stratigraphy: 713-762. Cambridge.
Galerkina, S. G. 1959. Stratigrafiya verkhnemelovykh u tretichnykh otlozhenii Berezovosko-Malo-
Atlymskogo raiona. [Stratigraphy of the upper Cretaceous and Tertiary deposits of the Berezov, Mal,
Atlym region.] Trudy vses. neft. nauchno-issled. geol.-razv. Inst. |VNIGRI] 140: 139-167.
Glezer, Z. I. 1966. Kremnevye Zhgutikovye Vodorosli (Silikoflagellaty). Flora Sporovykh Rastenii SSSR.
7: 331 pp. Moskva. Translation: Silicoflagellatophyceae. Cryptogamic Plants of the U.S.S.R. 7: 363 pp.
Jerusalem.
—— 1969. Pozdneeotsenovye Kompleksy Diatomovykh, kremnevykh zhgutikovykh vodoroslei i ebriidei
yugo-zapadnoi chasti Turgaiskogo Progiba. [Late Eocene complexes of diatoms, silicoflagellates and
Ebriidae of the south-western part of the Turgai Gates.] Trudy vses. nauchno-issled. geol. Inst. 130 (4):
67-85.
— 1975. K revizii roda Triceratium Ehr. sensu Hustedt, 1930 (Bacillariophyta). [Towards a revision of
the genus Triceratium Ehr. sensu Hustedt, 1930.] Bot. Zh. SSSR 60: 1304-1310, tab. I-III.
—— 1979. Evolyutsiya i sistematika poryadka Biddulphiales (Bacillariophyta). Paleont. Zh. 1979 (1):
110-120. Translation: Evolution and systematics of the order Biddulphiales (Bacillariophyta). Paleont.
J. 13: 103-111. ;
— 1983. Taksonomicheskaya znachimost’ priznakov u diatomovykh vodoroslei v svete razrabotki novoi
klassifikatsii Bacillariophyta. [Taxonomic significance of characters in diatoms in the light of elaboration
of a new Bacillariophyta classification.| Bot. Zh. SSSR 68: 993-1002.
— 1986. O polozhenii roda Triceratium s. str. (Bacillariophyta) v sisteme diatomovykh vodoroslei. [On
the position of the genus Triceratium s. str. (Bacillariophyta) in the diatom system.] Bot. Zh. SSSR 71:
1543-1545.
——, Jousé, A. P., Makarova, I. V., Proshkina-Lavrenko, A. I. & Sheshukova-Poretskaya, V. S. 1974.
Diatomovye Vodorosli SSSR Iskopaemye i Sovremennye. [The diatoms of the USSR fossil and recent. |
1. 403 pp. Leningrad.
Gombos, A. M., Jr. 1976. Paleogene and Neogene diatoms from the Falkland Plateau and Malvinas Outer
Basin. Initial Rep. deep Sea Drilling Proj. 36: 575—687.
— 1982. Early and middle Eocene diatom evolutionary events. Bacillaria 5: 225-242.
— 1983. Middle Eocene diatoms from the South Atlantic. Initial Rep. deep Sea Drilling Proj. 71:
565-581.
— 1984. Late Paleocene diatoms in the Cape basin. Initial Rep. deep Sea Drilling Proj. 73: 495-511.
—— & Ciesielski, P. F. 1983. Late Eocene to early Miocene diatoms from the southwest Atlantic. Jnitial
Rep. deep Sea Drilling Proj. 71: 583-634.
Gray, S. F. 1821. A natural arrangement of British plants 1. London.
Greville, R. K. 1863. Descriptions of new and rare diatoms. Series IX. Trans. microsc. Soc. Lond. II, 11:
63-76, pl. IV-V.
— 1865. Descriptions of new and rare diatoms. Series XIX. Trans. microsc. Soc. Lond. II, 13: 1-10, pl.
I-III.
294 R. ROSS AND P. A. SIMS
Grove, E. & Sturt, G. 1886. On a fossil marine diatomaceous deposit from Oamaru, Otago, New Zealand.
Part I. J. Quekett microsc. Club I, 2: 321-330, pl. XVII-XIX.
Grunow, A. 1884. Die Diatomeen von Franz Josefs-Land. Denkschr. Akad. Wiss. Wien 48 (2): 53-112,
Taf. I-V.
— 1887. [Review of Grove & Sturt, 1886.] Bot. Zbl. 31: 131-133.
Hajos, M. 1976. Upper Eocene and lower Oligocene Diatomaceae, Archaeomonadaceae and Silicoflagel-
latae in southwestern Pacific sediments, DSDP Leg 29. Initial Rep. deep Sea Drilling Proj. 35: 817-883.
Hasle, G. R., von Stosch, H. A. & Syvertsen, E. E. 1983. Cymatosiraceae, a new diatom family. Bacillaria
6: 9-156.
Heiberg, P. A. C. 1863. Conspectus criticus Diatomacearum Danicarum. 135 pp., 6 Taf. Kébenhavn.
Hendey, N. I. 1964. An introductory account of the smaller algae of British coastal waters. Part V:
Bacillariophyta (diatoms). Fishery Invest., Lond. IV, 5. pp. i-xxii, 1-317, pl. I-XLV.
Holmes, R. W. & Brigger, A. L. 1979. The marine fossil genus Entogonia Greville — a review. Bacillaria 2:
155-214.
Hustedt, F. 1930. Die Kieselalgen Deutschlands, Osterreichs und der Schweiz. In Dr L. Rabenhorst’s
Kryptogamen-Flora von Deutschland, Osterreich und der Schweiz 7 (1, 5): 785-920.
Jousé, A. P. 1977. Atlas Mikroorganizmov b donnykh Osadkakh Okeanov. Diatomei, Radiolyarii,
Silikoflyagellyaty, Kokkolity. Atlas of microorganisms in bottom sediments of the oceans. 32 pp., 160
Tabl. Moskva.
Kitton, F. 1870. Diatomaceous deposits from Jutland. J. Quekett microsc. Club 2: 99-102.
Kitzing, F. T. 1849. Species algarum. Lipsiae.
Laporte, L. J. & Lefébure, P. 1929. Diatomées rares et curieuses. 1 P|. 1—XV. Paris.
Lefébure, P. 1935. Diatomées contenues dans le dépét fossile marin de Saint-Laurent-la- Vernéde (Gard).
Bull. Soc. fr. Microsc. 4: 44—57, pl. WI-IV.
Miller, U. 1969. Fossil diatoms under the scanning electron microscope. A preliminary report. Sver. geol.
Unders. Afh. C, 63 (5): 1-65.
Miller, W. I. & Collier, A. 1978. Ultrastructure of the frustule of Triceratium favus (Bacillariophyceae). J.
Phycol. 14: 56-62.
Mukhina, V. V. 1974. Paleotsenovye Diatomovye ily v vostochnoi chasti Indiiskogo Okeana. [Paleocene
diatomaceous ooze in the eastern part of the Indian Ocean.] Okeanologiya 14: 852-858.
1976. Species composition of the late Paleocene diatoms and silicoflagellates in the Indian Ocean.
Micropaleontology 22: 151-158, pl. 1-2. ;
Navarro, J. N. 1981. Asurvey of the marine diatoms of Puerto Rico. II. Suborder Biddulphiineae: Families
Biddulphiaceae, Lithodesmiaceae and Eupodiscaceae. Botanica mar. 24: 615-630.
Pantocsek, J. 1889. Beitrdége zur Kenntniss der fossilen Bacillarien Ungarns. II Theil. 123 pp., 30 pl.
Nagy-Tapolcsany.
Paramonova, N. V. 1964. Materialy po diatomovym vodoroslyam iz paleogenovykh otlozhenii severa
zapadnoi Sibiri. [Materials on the diatoms from paleogene deposits of north-western Siberia.] Trudy
vses. nauchno-issled. geol.-razv. Inst. 239: 232-246, 476-497.
Proshkina-Lavrenko, A. I. (Ed.), Jousé, A. P. & Sheshukova, V. S. 1949a. Diatomovyi Analiz. Kn. 1.
Obshchaya i paleobotanicheskaya kharakteristika diatomovykh vodoroslei. [Diatom analysis. General
and paleobotanical characteristics of diatoms. ] 239 pp. Leningrad.
——.,, Kiselev, A. I., Poretskii, V. S. & Sheshukova, V. S. 1949b. Diatomovyi Analiz. Kn. 2. Opredelitel’
iskopaemykh i sovremennykh diatomovykh vodoroslei. Poryadki Centrales i Mediales. [Diatom
analysis. Descriptions of fossil and recent diatoms. Orders Centrales and Mediales.] 238 pp., 101 Tab.
Leningrad.
Ross, R. 1985. Proposal to conserve Hemiaulus Heiberg against Hemiaulus Ehrenberg (Bacillariophyta).
Taxon 34: 301-302.
——, Cox, E. J., Karayeva, N. I., Mann, D. G., Paddock, T. B. B., Simonsen, R. & Sims, P. A. 1979. An
amended terminology for the siliceous components of the diatom cell. Beih. nov. Hedwigia 64: 513-533.
& Sims, P. A. 1971. Generic limits in the Biddulphiaceae as indicated by the scanning electron
microscope. In V. H. Heywood (Ed.), Scanning electron microscopy, systematic and evolutionary
applications: 155-177. London.
— 1980. Syringidium Ehrenb., Dextradonator Ross & Sims, nov. gen. and Abas Ross & Sims, nov.
gen. Bacillaria 3: 115-127.
1985. Some genera of the Biddulphiaceae (diatoms) with interlocking linking spines. Bull. Br.
Mus. nat. Hist. (Bot.) 13: 277-381.
& Hasle, G. R. 1977. Observations on some species of the Hemiauloideae. Beih. nov. Hedwigia
54: 179-213.
GENERA OF THE BIDDULPHIACEAE 295
Round, F. E. 1981. The ecology of algae. 653 pp. Cambridge.
Rudkevich, M. Ya., Rubina, N. V. & Permyakov, A. I. 1957. Materialy k stratigrafii paleogena nizhnego
Priob’ya. [Materials towards the stratigraphy of the Paleogene of the lower Ob basin.] Byull. Mosk.
Obshch. Ispyt. Prir. 11, 62, Otdel Geol. 32 (1): 75-91.
Schmidt, A. 1890. Atlas der Diatomaceenkunde. Heft 38. Taf. 149-152. Leipzig.
—— 1893. Atlas der Diatomaceenkunde. Heft 46. Taf. 181-184. Leipzig.
Schrader, H.-J. & Fenner, J. 1976. Norwegian Sea Cenozoic diatom biostratigraphy and taxonomy. Initial
Rep. deep Sea Drilling Proj. 38: 921-1099.
Schulz, P. 1927. Diatomeen aus norddeutschen Basalttuffen und -Tuffgeschieben. Z. Geschiebeforsch.
3: 66-78, 113-126.
Schiitt, F. 1896. Bacillariaceae (Diatomeae). Jn A. Engler & K. Prantl, Naturl. PfiFam. 1 (1, b): 31-153.
Shibkova, K. G. 1968. Diatomei paleogenovykh otlozhenii Yuzhnogo Kazakhstana. [Diatoms of
Paleogene deposits of southern Kazakhstan.] Jn A. P. Jousé (Ed.), Iskopaemye Diatomovye Vodorosli
S.S.S.R.: 21-26. Moskva.
Shrubsole, W. H. & Kitton, F. 1881. The diatoms of the London Clay. J/ R. microsc. Soc. I, 1: 381-387.
Simonsen, R. 1974. The diatom plankton of the Indian Ocean Expedition of R/V ‘Meteor’ 1964-1965.
Meteor ForschErgebn. D, 19: 1-107.
—— 1979. The diatom system: ideas on phylogeny. Bacillaria 2: 9-71.
Sims, P. A. 1986. Sphynctolethus Hanna, Ailuretta gen. nov., and evolutionary trends within the
Hemiauloideae. Diatom Research 1: 241-269.
Smith, H. L. 1872. Conspectus of the families and genera of the Diatomaceae. II. Lens 1: 72-93.
Stosch, H. A. von 1985. Some marine diatoms from the Australian region, especially from Port Phillip Bay
and tropical north-eastern Australia. Brunia 8: 293-348.
—— & Simonsen, R. 1984. Biddulphiopsis, a new genus of the Biddulphiaceae. Bacillaria 7: 9-36.
Strel’nikova, N. I., Kaplan, A. A. & Travina, M. A. 1978. Paleogenovye diatomei, silikoflyagellyaty i
ebriidei Kaliningradskoi oblasti. [Paleogene diatoms, silicoflagellates and ebriidei of Kaliningrad
oblast.] Jn A. P. Jousé (Ed.), Morskaya Mikropaleontologiya (Diatomei, Radiolyarii, Silikoflyagellyaty,
Foraminifery i izvestkovyi Nannoplankton): 57-66.
Tempére, J. & Peragallo, H. 1889a. Diatomées, Fasc. 1. pp. 1-16. Paris.
—— — 1889b. Diatomées. Fasc. 2. pp. 17-32.
—— —— 1890. Diatomées. Fasc. 7. pp. 97-112.
—— — 1891. Diatomées. Fasc. 15. pp. 225-240.
—— — 1907. Diatomées du monde entier. Fasc. 1. pp. 1-16. Paris.
—— —— 1908. Diatomées du monde entier. Fasc. 5. pp. 65-80.
— —— 1911. Diatomées du monde entier. Fasc. 11. pp. 161-176.
—— —— 1912. Diatomées du monde entier. Fasc. 22. pp. 337-352.
— —— 1913. Diatomées du monde entier. Fasc. 27. pp. 417-432.
Truan y Luard, A. & Witt, O. N. 1888. Die Diatomaceen der Polycistinenkreide von Jérémie in Hayti,
Westindien. Berlin.
Tsumura, K. 1964. Some kinds of Rutilaria and Pseudorutilaria. Bull. Yokohama City Univ. Soc. 16:
77-96, Pl. I-III.
Van Heurck, H. 1883. Synopsis des diatomées de Belgique, Atlas, fasc. 6, pl. 104-132, 22 bis, 82 bis, 83 bis,
83 ter, 95 bis. Anvers.
—— 1896. A treatise on the Diatomaceae, translated by Wynne E. Baxter. xx + 558 pp., 35 Pl. London.
Vozzhennikova, T. F. 1960. Paleoalgologicheskaya kharakteristika mezokainozoiskikh otlozhenii zapad-
no-sibirskoi nizmennosti. [Paleoalgological characteristics of the mesocaenozoic deposits of the West-
Siberian plain.] Trudy Inst. Geol. Geofiz. sib. Otd. 1: 7-64.
Walker, W. C. & Chase, H. H. 1886. Notes on some new and rare diatoms. (Series I). 7 pp., 2 Pl. Utica, New
York.
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 Jousé, Northern Urals, Tyumen’sk
oblast, U.S.S.R.; Figs 1, 3, 4: bar line = 10 wm; Figs 2, 5, 6: bar line = 2 wm.
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 wm; Figs 10, 12, 13, 14: bar line = 3 wm.
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° 08’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 line = 20 wm; Figs 18, 21—23: bar line = 5 wm.
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 wm; Figs 29-33, bar line = 5
pm.
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: linked 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 wm; Figs 36—38: bar line = 5 wm.
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 um; Figs 42-44: bar line = 2 um; Figs 45—47: bar line = 5 wm.
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° 01'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, 1525m
depth, Vema cruise 17, core 107, 50 cm. Figs 48, 50: bar line = 20 wm; Figs 49, 51-54: bar line = 5 wm.
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-Vernéde, Gard, France. Figs 55—60: bar line
= 25 wm; Figs 61-64: bar line = 5 wm.
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 um; Figs 66-70; bar line = 5 wm.
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 simulans R. Ross & P. A. Sims. ‘Carlovo’, U.S.S.R. Figs 71, 72, 74: bar line = 20 wm; Figs
73, 75-77: bar line = 3 wm.
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 wm; Figs 84-85: bar line = 10 wm.
Identity of specimen shown in Figs 82—3 doubtful (see p. 289).
Fig. 78: two valves linked, upper with hyaline band (pleura) attached. Note linking mechanism involving
two flattened and acute linking 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: linked valves, girdle view. Note two spines from each valve
projecting beyond the girdle bands. Fig. 83: oblique view, valve face with small hyaline central area,
radiating rows of fine poroid areolae extending to marginal ridge and slightly 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
hyaline 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 um. Fig.
87: fo. pentagona Jousé, BM 65833: 36 wm diameter. Fig. 88: fo. exsculptum BM coll. Adams TS748: full
length one side, 67 wm. 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 wm, b: 97 wm, c: 130 wm. 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
pm; Fig. 91: Holotype; apical axis 95 wm. 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 wm. 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 wm.
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
pum. Fig. 95: two linked valves; apical axis 80 wm. Fig. 96: Pseudorutilaria hendeyi R. Ross & 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 wm. Figs 97-98: Monile laurentii R. Ross & P. A. Sims, St
Laurent-la-Vernéde, Gard, France. Fig. 97: Holotype, BM coll. Adams H941, specim. I, 12: apical axis
220 um. Fig. 98: BM 81303: apical axis 228 wm. Figs 99-100: Bonea simulans R. Ross & P. A. Sims,
‘Carlovo’, U.S.S.R.; Fig. 99: Holotype, BM 81301: length of side 119 um. Fig. 100: BM 81302: girdle view;
length of side 140 um.
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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.
0 565 00777 7 £30.00
Vol. 2. The Dicotyledons
(Part). 1979, 220pp.
0 565 00810 2 £30.00
Vol. 3. The Dicotyledons
(Part). 1982, 226pp, 1 fig.
0 565 00854 4 £35.00
Titles to be published in Volume 16
Studies in the genus Hypericum L. (Guttiferae)
7. Section 29. Brathys (part 1)
By N. 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
Further genera of the Biddulphiaceae (diatoms) with interlocking linking spines
By R. Ross & P. A. Sims
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd, Dorchester
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