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BULLETIN OF
THE BRITISH MUSEUM
(NATURAL HISTORY)

BO LANY:
VOL. 3

1963—65

TRUS DEES. Or

THE BRITISH MUSEUM (NATURAL HISTORY)

LONDON: 1965

DATES OF PUBLICATION OF THE PARTS

No. I, May 1963
No. 2, July 1963
No. 3, September 1963
No. 4, September 1964
No. 5, May 1965
No. 6 October 1965

Contents and Index, October 1965

PRINTED IN
GREAT BRITAIN
AT THE
BARTHOLOMEW PRESS
DORKING
BY
ADLARD AND SON, LTD.

CONTENTS

BOTANY VOLUME 3

PAGE
A revision of the genera Buchenavia and Ramatuella. By A. W.
EXELL and C. A. STACE >
The diatom genus Capartogramma and the identity of Schizostauron.
By R. Ross 47
Angiosperms of the Cambridge Annobon Island Expedition. By
A. W. EXELL 93
A revision of the genus Petrorhagia. By P. W. Baty and V. H.
HEYWoopD I19Q
Marine algae of Gough Island. By YVoNNE M. CHAMBERLAIN 173
The Ceylon species of Asplenium. By W. A. SLEDGE 233

Index to Volume 3 270

“¢! GS ati +
uF, Geis i 2 FE
+ | atang Maier 4

es |) ae, ae : fp
os ns ae ae se
tna ot So eae
peat eo : ae
Renta. a) a
rye mn
ent ae i
ay 03 Weis

OE ae ar peri age =

A REVISION OF THE GENERA
BUCHENAVIA AND
RAMATUELLA

A. W. EXELL

AND

GA. 8 FACE

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

_ BOTANY | Vol. 3 No. 1
Fig LONDON: 1963 ©

a REVISION OF THE GENERA
BUCHENAVIA AND RAMATUELLA

Bivs

AL We EXELL ann: CoA. STACE
ways

Pp. 1-46 ; 5 Text-figures

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

BOTANY Vol. 3 No. 1
LONDON : 1963

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), tstituted in 1949, 1s
issued in five series corresponding to the Departments
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This paper is Vol. 3, No. 1 of the Botany series.

© Trustees of the British Museum, 1963

PRINTED BY ORDER OF THE TRUSTEES OF
THES BRIEISH MUSEUM

Issued May 1963 Price Sixteen Shillings

A REVISION OF THE GENERA
BUCHENAVIA AND RAMATUELLA

By A. W. EXELL and C. A. STACE!

THE two genera Buchenavia Eichl. and Ramatuella Kunth are confined to tropical
America, in common with two other genera of the family Combretaceae (Bucida and
Thiloa). While one of us (C. A. S.) was undertaking a detailed study of the cuti-
cular anatomy of Buchenavia and Ramatuella, as part of a comprehensive survey
of the whole family, it became apparent that the taxonomy of the former genus was
badly in need of revision. Although twenty-eight specific names had appeared
under Buchenavia it had never been revised. Since certain changes in the taxonomy
of Ramatuella also seemed advisable, a joint revision of both genera was undertaken.

As many of the earlier, and some of the later, species of Buchenavia and Ramatuella
have been very inadequately described, and the discovery of new species has revealed
further characters of diagnostic significance, descriptions of all the species of these
two genera are included in this paper.

Our work is based mainly on the collections in the herbaria of the British Museum
(Natural History) (BM) and the Royal Botanic Gardens, Kew (K). Loans were
also received from the Chicago Natural History Museum (F), Botanical Institute
of the Academy of Sciences of the U.S.S.R., Leningrad (LE), Botanische Staats-
sammlung, Munich (M), New York Botanical Garden (NY), Muséum National
d’Histoire Naturelle, Laboratoire de Phanérogamie, Paris (P), U.S. National Museum,
Washington (US), and Naturhistorisches Museum, Vienna (W). We are grateful
to the Directors of these Institutions, and especially to the Botanical Institute of
the Academy of Sciences of the U.S.S.R., and to Dr. J. Cuatrecasas of Washington,
for the donation to the British Museum of duplicate specimens. Concurrently
with the commencement of this revision a fairly large and very useful collection
of the two genera concerned was received at the British Museum from the Instituto
Agrondémico do Norte, Belém, Brazil (IAN).

The abbreviations indicated parenthetically above are used in the citation of
specimens throughout the paper. Other institutions cited are: Natuurweten-
schappelijk Museum, Antwerp (AWH) ; Botanisches Museum, Berlin (B) ; Jardin
Botanique de l’Etat, Brussels (BR) ; Museu Paraense Emilio Goeldi, Belém (MG) ;
Jardim Botanico, Rio de Janeiro (RB); Botanical Department, Naturhistoriska
Riksmuseum, Stockholm (S).

1 During this investigation C. A. S. was receiving a grant from the Department of Scientific and
Industrial Research, for which he is most grateful.

BOT. 3, I. 1§

4 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA
BUCHENAVIA Eichl.

BucHENAVIA Eichl. in Flora 49:164 (1866); in Mart., Fl. Brasil. 14 (2): 95
(1867).
Type: B. capitata (Vahl) Eichl.

Buchenavia belongs to the tribe Terminalieae of the family Combretaceae and is
most closely related to Terminalia, from which it differs in that the filaments of the
stamens are adnate to the back of the anthers and the calyx lobes are only very
slightly or scarcely at all developed. In Tevminalia the anthers are versatile and
the calyx lobes well developed, usually triangular in shape and acute at the apex.

As in all genera of the Combretaceae except Strephonema, the “ receptacle ”’ of
Buchenavia is composed of two morphological components: an upper portion
(“upper receptacle’ or “ calyx tube ’’), cup-shaped in this genus (sometimes con-
siderably elongated in Combretum and Quisqualis), and a lower portion (‘‘ lower
receptacle ’’) surrounding and fused with the ovary and (in this genus) extended
above it. As it is usually not easy or possible to ascertain from gross morpho-
logical studies which part of the “‘ upper receptacle ”’ is in fact receptacular rather
than appendicular (although in Laguncularia and Lumnitzera the presence of adnate
bracteoles indicates the probable receptacular origin of at least part of that organ),
we have used the term “ hypanthium ”’ rather than “ receptacle”’. In Buchenavia
the cupuliform upper hypanthium is almost always broader than long and bears
five scarcely developed calyx lobes, ten stamens, no petals, and a well-developed
hairy disk. It is usually more elongate (i.e. campanulate) in Terminalia. The
lower hypanthium is also characteristic in Buchenavia, consisting of a proximal
portion surrounding the sessile ovary and a distal portion tapering from the apex
of the ovary and simulating a pedicel, bearing the upper hypanthium at its apex.
In most species the portion surrounding the ovary is pubescent, while the distal
portion and the whole of the outside of the upper hypanthium are glabrous or almost
so. In one or two species, however, the lower portion is also glabrous. In
Terminalia the whole hypanthium is usually uniformly hairy or (less often) glab-
rous, but a few species have the Buchenavia pattern of indumentum. Some species
of Terminalia possess a lower hypanthium shaped as in Buchenavia, but the
majority lack the long pedicel-like distal portion.

Eichler (1866, 1867) included eight species in the genus: one (B. capitata (Vahl)
Eichl.) previously known in Bucida and also in Terminaha; one (B. oxycarpa
(Mart.) Eichl.) previously known in Terminalia ; and the other six newly described.
Since then twenty additional names have appeared in the literature (three of these
being nomina nuda, the rest validly published) but the genus has not been revised.
Out of the 25 valid names we recognize 21 species, and we also describe three addi-
tional ones. From various herbarium specimens we have seen it seems that there
may be several more awaiting description but they are at present insufficiently
known.

The genus is confined to but fairly widespread in tropical America where it is
centred on the Amazon basin (see Figs. 1, 2). It occurs throughout the West Indies
(one species) from Cuba (within 150 kilometres south of the Tropic of Cancer) to

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 5

Trinidad ; in Panama (one species); Venezuela (five species) ; Colombia (five
species) ; the Guianas (four species) ; Brazil (twenty species) ; Peru (two species) ;
and Bolivia (one species). One outlying species (B. kleinii) occurs 500 kilometres

Tropic of Cancer

V
— °9 aS
°
Soe 9
_ a ee

Puerto Rico
AS Laer.
Jamaica S——_—— Hispaniola ~— S.__ o ¢ Antigua
Montserrat °@ Guadeloupe
Dominica,°
© Martinique

0
St. Vincent ° \o

Panama

Equator

ee BRAZIL

Tropic of Capricorn

Fic. 1. Distribution of Buchenavia (part) and Ramatuella: 1, B. capitata (insular) ;
2, B. parvifolia; 3, B. kleinii; 4, B. ochroprumna; 5, Ramatuella (all species) ; C,
B. capitata (continental) ; S, B. sericocarpa,

6 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

Equator

Tropic of Capricorn

Fic. 2. Distribution of Buchenavia (part): 1, B. reticulata, B. suaveolens and B. ptero-
carpa; 2, B. tomentosa; 3, B. oxycarpa; 4, B. macrophylla; 5, B. fanshawei; 6,
B. grandis; Ac, B. acuminata; Ca, B. callistachya; Co, B. congesta; D, B. discolor ;
H, B. huberi; L, B. longibracteata; Me, B. megalophylia; O, B. oxycarpa (outlier) ;
Pa, B. pallidovirens ; Pl, B. pulcherrima; Pn, B. punctata; R, B. reticulata (outlier) ;
V, B. viridifilora.

south of the Tropic of Capricorn in southern Brazil. This distribution pattern
follows that of the family as a whole (in the New World) and is a typical pattern
for a predominantly tropical group. The largest two genera, Combretum and
Terminalia, are almost pantropical; the two small genera Laguncularnia and
Conocarpus are found on both sides of the Atlantic ; Buchenavia is the largest genus
of Combretaceae confined to one hemisphere, this being due no doubt to the well-
known phenomenon of prolific speciation in the Amazonian forest region ; Bucida,
Ramatuella and Thiloa, the other three American genera, are also confined to the
New World.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 7

Eichler (1866) recognized two groups of species in Buchenavia : “‘ Flores fructusque
capitati’’ and “ Flores fructusque vel saltem fructus spicati’’. It is still possible
to recognize these two groups with ease, and we have used this character in the key.
We have refrained, however, from giving the two groups taxonomic status, since
there appear to be no other constant differences between them, and from a study
of other genera in the Combretaceae (especially Combretum) it seems that the capitate
inflorescence has arisen on several separate occasions in rather distantly related
groups. Furthermore, the capitate B. sericocarpa is otherwise almost identical
(as far as we can tell) with the spicate B. acuminata.

In the flowering state it is usually very easy to distinguish between species of
the two groups, as the inflorescences of one are long and slender, the flowers in many
cases being well separated from each other, while in the other group the flowers
are crowded together on a rhachis rarely as long as the peduncle. In the fruiting
state the distinction is no more difficult, although usually very few fruits reach
maturity in each infructescence. The presence of a spicate inflorescence is, however,
always evident, even when the peduncle bears but a single fruit, since the scars
of the fallen flowers may be seen at intervals along the rhachis. In this spicate
group the rhachis often elongates considerably during ripening of the fruits.

Much of the difficulty encountered in the genus is due to the incompleteness of
our knowledge of several species which are known from fruits and leaves or from
flowers and leaves only. The leaves are frequently of a very different appearance
at flowering and fruiting stages. Fortunately, although the fruits of Buchenavia
do not differ from those of Terminalia as a whole, all the American species of the
latter genus apparently have conspicuously winged fruits, while all species of
Buchenavia have fruits without wings or with very narrow ones. The greatest
development of wings is shown by B. pterocarpa and B. punctata. In using this
character to separate the two genera, the only possible source of error would be
in the case of Terminalia catabpa, now widespread in tropical America, but this is
an introduced Asiatic and Australasian species easily recognizable by its large thin
obovate leaves usually with a cordate or subcordate base.

Buchenavia species are shrubs or trees from about three to fifty metres tall.
Ecologically they may apparently be roughly divided into two main groups: small
trees or shrubs of lowland forest often inundated by water, frequently growing on
river banks, by lakes and in swamps; and tall trees of more upland rain forest.
The latter are said to flower “‘ very seldom” (Ducke in Bol. Técn. Inst. Agron.

Norte 4 : 25 (1945)).

KEY TO THE SPECIES

In the following key we have used the characters which are most likely to be
present on herbarium specimens: thus, unless absolutely necessary, fruit characters
alone are not used. This has sometimes resulted in long couplets citing exceptions
but is thought to be justified by the added usefulness in identifying incomplete
specimens. The flowers unfortunately appear to be of very little taxonomic help in
this genus. As in the descriptions, the terms “ tomentellous’”’ (a diminutive of
“tomentose ”’) and “puberulous ’”’ (a diminutive ot “ pubescent ’’) are taken to

BOT. 3, I. 1§§

8 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

indicate not a lesser amount of indumentum but a shorter length of the hairs.
‘‘ Pseudostipe ”’ indicates a stalk-like basal portion of the fruit, a true pedicel being
absent. Leaf length always includes the petiole (except, of course, when leaf-
lamina is specified). In cases where the lamina is decurrent into the petiole it is
not feasible to determine an exact point at which to distinguish between them.
We have occasionally referred to the indumentum (or lack of indumentum) of the
‘ovary ’’, this being merely a short way of saying “‘ that part of the lower hypan-
thium enclosing the ovary”’. This part usually differs in indumentum from the
apical narrowed part.

Flowers in short ovoid or capitate spikes usually less than 1-5 cm. long; peduncle
usually longer than the rhachis ; spikes not elongating in fruit :

Fruit glabrous or rarely very sparsely pubescent :

Leaf-lamina coriaceous when mature, c. 2-8 X I-4 cm., narrowly to broadly
obovate ; major lateral nerves rather conspicuous, c. 2-6 pairs; petiole
c. 4-20 mm. long, usually rather stout ; peduncle slender or stout ; flowers
with densely pubescent ovary; fruit 18-30 x Io-20 mm.; endocarp
markedly longitudinally ridged ; smallest twigs rather stout (widespread
throughout tropical America from Cuba to S. Brazil) . 1. B. capitata

Leaf-lamina chartaceous when mature, c. I-5-4 X 0*5-2cm., narrowly to broadly
obovate ; major lateral nerves rather inconspicuous, c. 5-12 pairs ; petiole
c. 2-5 mm. long, slender ; peduncle slender ; flowers with glabrous ovary ;
fruit 14-20 X 8-12 mm.; endocarp indistinctly longitudinally ridged ;
smallest twigs very slender (Amazon and Orinoco basins only)

2. B. parvifolia

Fruit pubescent or puberulous :

Fruit densely but minutely puberulous, obtuse or acute at the apex; leaves
obovate, usually broadly so, rounded, obtuse or rarely very shortly api-
culate at the apex (S. Brazil only) : ; ; 3. B. klein

Fruit densely tomentose ; leaves narrowly to broadly obovate, rounded to long-
acuminate at the apex (N. Brazil only) :

Fruit silvery- or grey-tomentose, obtuse at the apex, smooth-surfaced ;
leaf-lamina oblong-obovate, 5-12 cm. long, apex abruptly caudate-
acuminate with an acute acumen ; lateral nerves 7-9 pairs

4. B. sericocarpa

Fruit ferrugineous-tomentose, long-beaked, markedly and irregularly ridged ;
leaf-lamina obovate to oblanceolate, 4-8 (10) cm. long, apex rounded ;
lateral nerves 3-5 pairs . : $ ; 5. B. ochroprumna

Flowers in long narrow spikes usually 4 cm. iene or more, if in shorter spikes then
the latter elongating in fruit :

Leaf-lamina chartaceous, up to 15 cm. long, very narrowly obovate to oblanceolate,
subacute to acute at the apex, conspicuously but minutely pellucid-punctate ;
fruit very strongly and symmetrically 5 (6)-ridged, apiculate to shortly
(to 3 mm.) acuminate at the apex, 27-31 mm. long . , 6. B. punctata

Leaf-lamina various, but never pellucid-punctate ; fruit usually more or less

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 9

terete or obscurely ridged, or if strongly ridged then 2 ridges much more

developed than the others or apiculus 3-9 mm. long :

Leaves at anthesis and usually for some time afterwards densely rubiginous-
to ferrugineous-pubescent to -tomentose below, pubescent and nearly
always drying reddish-brown above; reticulation usually very con-
spicuously raised below at maturity, connecting the 5-15 pairs of lateral
nerves :

Leaf-lamina (2) 5-26 (36) < I-7 (12) cm., narrowly elliptic to oblanceolate,
obtuse to acute or shortly apiculate at the apex ; inflorescence 7-18 cm.
long ; fruit ferrugineous- or rubiginous-tomentose or densely pubescent,
apiculate to acuminate at the apex . : ; . 7. B. reticulata

Leaf-lamina 3:°5-13°5 X I°5-7 cm., elliptic or obovate (narrowly elliptic or
oblanceolate when very young), rounded at the apex (obtuse or more
pointed when very young) ; inflorescence 25-13 cm. long; fruit sparsely
ferrugineous-pubescent, rounded at the apex :

Leaf-lamina rather conspicuously reticulate below when mature ; inflores-
cence 3°5-I13 cm. long; flowers c. 4-5 X 2°5-4 mm.; fruit 22-31
x 15-25 mm., rather to very sparsely puberulous 8. B. tomentosa

Leaf-lamina very conspicuously reticulate below when mature ; inflores-
cence 2:5-5°5 cm. long; flowers c. 2:5-3 X I°75-3 mm. ; fruit 21-23
< 12-15 mm., puberulous . ; : an E B. callistachya

Leaves becoming sparsely pubescent to almost ginbroue some time before an-
thesis, rarely drying reddish-brown above ; reticulation never conspicuously
raised (though lateral nerves often so) :

Leaves almost or quite glabrous, up to 11-5 cm. long but often much shorter ;
lateral nerves inconspicuous below, scarcely or not at all raised, domatia
entirely absent from their axils; fruit flattened or terete, glabrous,
with a distinct apiculus up to 4 mm. long :

Leaf-lamina narrowly elliptic or oblanceolate to elliptic or obovate (usually
obovate) ; fruit 24-30 mm. long, more or less terete, apiculus usually
straight ; endocarp smooth, almost round in transverse section

10. B. suaveolens

Leaf-lamina narrowly elliptic or oblanceolate to narrowly obovate ; fruit
14-28 mm. long, flattened with two narrow lateral wings and three
additional] facial ridges, apiculus usually curved ; endocarp irregularly
furrowed, with two lateral wings : . 11. B. pterocarpa

Leaves various ; lateral nerves conspicuous below, the major nerves consider-
ably or somewhat raised and very often with domatia in some or all of
their axils ; fruit various, often long-beaked or rounded at the apex :

Inflorescence with conspicuous bracts c. 7-13 mm. long subtending and
exceeding the flowers; bracts persistent at least until anthesis or
after; leaf-lamina 8-23 cm. long, oblanceolate, acuminate at the
apex ; petiole up to 1:5 cm. long . . . 20. B. longibracteata

Inflorescence with small narrow bracts subtending the flowers, bracts
rarely equalling the length of the open flowers, up to c. 5 mm. long,

io REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

usually caducous before anthesis; (flowering material unknown in

B. pulcherrima, but this with petiole over 3 cm. long) :

Fruit with an apical beak (3) 5-20 mm. long and abruptly constricted
from the rest of the fruit :

Fruit scarcely or not pseudostipitate at the base, beak up to 13 mm.
long ; leaf-lamina up to 13 X 5 cm., lateral nerves 3-9 (12) pairs :

Leaves with (4) 6-9 (12) pairs of lateral nerves ; fruit quite glabrous
12. B. oxycarpa

Leaves with 3-4 pairs of lateral nerves ; fruit tomentellous
13. B. discolor

Fruit with a pseudostipe 2-7 mm. long, beak 10-20 mm. long ; leaf-

lamina c. (15) 20-32 X 5-10 cm., lateral nerves (g) 10-14 pairs
14. B. megalophylla
Fruit rounded to acute or apiculate at the apex, apiculus when present
very short (abnormally in young B. fanshawet apiculus up to 2 mm.
long) :

Leaf-lamina up to II X 4 cm., oblanceolate, abruptly and acutely
apiculate or mucronate or shortly acuminate at the apex ; lateral
nerves 5-7 pairs, arising at about 40° or less and then curving
towards the leaf-apex at about 25° or less for most of their length ;
reticulation conspicuous, very slightly raised below with very
small inter-reticular spaces ; fruit glabrous, ellipsoid-obovoid

15. B. pallidovirens

Leaf-lamina various, often much larger than Ir x 4 cm., if smaller
rarely acutely apiculate or acuminate at the apex; lateral nerves
3-—numerous pairs, very rarely or never less than about 40° to the
midrib for their proximal half ; fruit various, often pubescent :

Leaf-lamina up to 10 X 5°5 cm. but rarely over 8 X 4 cm.; apex
acute to rounded or retuse, never apiculate or acuminate ;
lateral nerves 4-7 pairs; inflorescence 2~7 cm. long; fruit
densely greyish-brown-tomentellous; domatia never as in
B. fanshawet (q. v.) : ; . 16. B. viridiflora

Leaf-lamina usually in all or most leaves over 8 cm. long, if less then
apex apiculate or acuminate, except in B. fanshawei (with
distinctive domatia, q.v.) and B. grandis (with glabrous fruit
and ovary often glabrous) :

Leaf-lamina up to 9°5 x 5 cm., elliptic or narrowly elliptic, apex
obtusely or acutely acuminate ; domatia never as in B. fan-
shawei (q.v.) ; fruit densely silvery-brown-tomentellous

17. B. acuminata

Leaf-lamina in all or most leaves usually I0 cm. or more long,
if less then apex not acuminate (but apiculate to rounded or
retuse) except in some forms of B. macrophylla (with less
hairy brownish fruit) and of B. fanshawei (with fruit similar
to the latter and distinctive domatia) :

REVISION @®F THE GENERA BUCHENAVIA AND RAMATUELLA 11

Leaf-lamina up to 14 X 6 cm., rounded to apiculate (acuminate)
at the apex ; lateral nerves 3-5 pairs; petiole sometimes
but usually not biglandular near the apex; distinctive
domatia present in nerve axils on lower surface of lamina
on some or most leaves of older or sometimes young branch-
lets, opening of domatium narrower than its diameter ;
fruit rather densely but minutely tomentellous

18. B. fanshawer

Leaf-lamina various, often over 15 cm. long ; lateral nerves (4)
6-c. 14 pairs, if 5 or less then leaf apex apiculate or acumin-
ate and petiole biglandular ; domatia absent or present
and then simple and triangular, pocket-like, with the
opening as wide as the diameter of the domatium ; fruit
various :

Petiole conspicuously biglandular in most Jeaves; fruit
minutely but rather densely brownish-tomentellous
19. B. macrophylla
Petiole without glands or petiolar glands inconspicuous ;
fruit either glabrous or densely greyish-brown- or rufous-
pubescent :

Leaf-lamina 27-29 x 8-10 cm., apiculate at the apex ; petiole
3.cm. or more long ; inflorescence c. 10-15 cm. long ;
fruit rounded to obtuse at both ends, densely rufous-
velutinous (British Guiana only) 21. B. pulcherrima

Leaf-lamina not over 16 cm. long; petiole up to about
3:2 cm. long; fruit glabrous or pubescent and then
apiculate at the apex, usually shortly pseudostipitate
at the base when mature (N. Brazil only) :

Fruit densely pubescent or tomentose at maturity,
indumentum wearing off when fruit very old but
still visible in parts . : . 22. B. huben

Fruit glabrous from an early stage; ovary sometimes
also glabrous :

Leaves not very densely clustered at branchlet tips,
up to 14 cm. long; lateral nerves 6-9 pairs ;
petiole up to 2 cm. long ; fruit with a pseudostipe
c. 2-3 mm. long, acutely apiculate at the apex

23. B. grandis

Leaves very densely clustered at the branchlet tips,
to-18 cm. long; lateral nerves g—I3 pairs;
petiole 2-3 cm. long; fruit with a pseudostipe
under 2 mm. long or not pseudostipitate, rounded
to more or less obtuse at the apex

24. B. congesta

12 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

1. Buchenavia capitata (Vahl) Eich]. in Flora 49: 165 (1866); in Mart., FI.
Brasil. 14 (2) : 96 (1867).

Bucida capitata Vahl, Eclog. Amer. 1: 50, t. 8 (1796).

Terminalia obovata Cambess. in St.-Hil., Fl. Brasil. Merid. 2 : 241 (1829).

Terminalia hilariana Steud., Nomencl. Bot., ed. 2, 2 : 668 (1841), nom. illegit.

Bucida angustifolia Spruce ex Eichl., loc. cit. (1867), nom. syn.; non B. angustifolia DC.
(1828).

Buchenavia vaupesana Cuatrec. in Fieldiana, Bot. 27 (1) : 108 (1950).

Buchenavia ptariensis Steyerm. in Fieldiana, Bot. 28 : 423 (1952).

Leaf-lamina membranous when young, coriaceous when mature, very variable
in size, narrowly to broadly obovate, retuse or rounded at the apex, sometimes
obtuse or subacute when very young, cuneate and sometimes narrowed at the base,
decurrent into the petiole, densely pubescent when very young but becoming glabrous
except on the midrib and lateral nerves when mature, often ciliate at flowering time ;
midrib and major lateral nerves usually conspicuous below, the latter variable in
number, 2-6 pairs; reticulation usually distinct when mature, slightly raised
below ; petiole 4-20 mm. long, usually rather stout. Peduncle slender or stout, c.
6-23 mm. long, almost glabrous to rufous-pubescent when young, almost glabrous
at maturity ; rhachis under 10 mm. long, often only 3 mm. long ; flowers in sparse
or dense capitula, Io-many in each inflorescence ; bracts very small and caducous
before anthesis. Upper hypanthium 1-1-5 mm. long, 2-3 mm. across, glabrous or
sparsely pubescent outside, pubescent within and ciliate on the margin; lower
hypanthium pedicel-like with the ovary at the base, 1-5-3 mm. long, densely rufous-
sericeous. Stamens exserted for c. 0-5-1°5 mm. Fruit ellipsoid, 5-ridged to sub-
terete, 18-30 X IO-20 mm., apiculate or acute to broadly obtuse at the apex, nar-
rowed below into a very short pseudostipe, glabrous or rarely very sparsely pubescent,
apparently rarely more than one ripening in each infructescence ; endocarp markedly
longitudinally ridged, usually about five ridges more prominent than the rest.

B. capitata is by far the most variable, abundant and widely distributed species
in the genus. It is found in the West Indies from Cuba (within 150 kilometres south
of the Tropic of Cancer) to Trinidad ; Panama; Colombia (Vaupés) ; Venezuela
(Amazonas and Bolivar) ; Dutch and French Guiana ; Brazil (Amazonas, Maranhao,
Ceara, Pernambuco and Rio de Janeiro within 150 kilometres north of the Tropic
of Capricorn) ; and Bolivia (Santa Cruz and La Paz). It is the only Buchenavia
found in the West Indies, where it occurs as a common tree of river banks and low-
land forest. It also appears to be frequent in similar conditions in north-east South
America, becoming rare westwards in Colombia and Venezuela and southwards in
southern Brazil and Bolivia (Fig. 1). Glaziou says (in Mém. Soc. Bot. France r (3) :
203 (1908)) that it is cultivated in Rio de Janeiro.

As it is a well-known species, described in many Floras, we have not cited the
numerous specimens preserved in herbaria.

We consider that B. ptariensis Steyerm. and B. vaupesana Cuatrec. fall within the
wide range of variation shown by B. capitata. In the West Indies plants of B. cap1-
tata are characterized by the large number of flowers in each inflorescence, the rather
small flower size and the relatively stout peduncles. Plants from the Amazon basin

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 13

tend to have fewer rather large flowers in each inflorescence and often more slender
and more glabrous peduncles. Specimens from coastal areas of the mainland are
often more or less intermediate in these characters. We are of the opinion that, in
the present state of our knowledge, this variation does not lend itself to subdivision
of the species into lower categories, but that it represents an ecocline from the
West Indies through the coastal areas of the South American continent to the
Amazon basin.

B. vaupesana appears to be an extreme example of the Amazon type, with rather
few large flowers and slender peduncles. We have seen the other characters by which
this ‘‘ species’ is said to differ from B. capitata (narrow somewhat pointed leaves
and long petioles) in other specimens of B. capitata, mostly due to immaturity.
The single known gathering (Cuatrecasas 6828), of which we have seen an isotype
(US), is the only record we have of B. capitata from Colombia. It is in flower.

B. ptariensis, of which we have seen the only known gathering, Steyermark
60038 (F, holotype), is a very small-leaved plant lacking flowers and fruit. We have
no evidence that it actually has a capitate inflorescence but it appears to fall at one
extreme of the B. capitata range of variation and we can see no reason for keeping it
separate. The leaves are of quite a different texture, shape and venation from those
of B. parvifolia, as the author points out. It is one of only two specimens of
B. capitata that we know of from Venezuela.

It seems quite possible that at some future date, when the material of B. capitata
is really representative throughout its wide range, it will become advisable to divide
the species into infraspecific units.

Vahl, when originally describing Bucida capttata, cited specimens collected by Ryan
in Montserrat which, after being described by Vahl, were sent to Banks. They are
now in the British Museum Herbarium (W. R. Dawson, Banks Letters : 727 (1958) ;
letter of r May 1794). There are two specimens, one with flowers, young leaves and
old fruits corresponding well with Vahl’s t. 8 [fig. 1], and the other with adult leaves
and extremely young inflorescences fitting his fig. 2. Since Vahl (tom. cit. : 51)
refers to the latter as ‘“‘ Aliam quoque in Montserrat legit Ryan ”’ and later (op. cit.
3 : 55 (1807)) indexes it as “ varietas ? ’’ it is reasonable to regard the former specimen,
with flowers and old fruits, as the holotype of B. capitata.

The vernacular name is given as “ Tanibuca’”’ by Glaziou (in Mém. Soc. Bot.
France I (3) :203), and Exell (Fl. Suriname. 3 (1) :175 (1935)) quotes “‘ Katoelima”’,
“ Toekoeli’’, ‘‘ Toekadi’’, “‘Matakki’’ and ‘‘Gemberhout ”’.

2. Buchenavia parvifolia Ducke in Arch. Jard. Bot. Rio Janeiro 4 : 150 (1925).

Leaf-lamina chartaceous when mature, rather small, c. 1-5-4 0:5—2 cm., narrowly
obovate to obovate, usually rounded but sometimes slightly retuse or very broadly
obtuse at the apex, cuneate at the base, narrowed and decurrent into the petiole,
almost glabrous except for sparse pubescence on the midrib and lateral nerves when
mature ; major lateral nerves often not very distinct, rather numerous for small size
of leaf and often with many minor lateral nerves almost as conspicuous, c. 5—I2 pairs ;

14 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

petiole slender and short, c. 2-5 mm. long. Inflorescences (fide Ducke) more or less
as in B. capitata ; peduncle c. 10-18 mm. long, slender. Ovary completely glabrous.
Fruit ellipsoid or less often obovoid, more or less terete, sulcate when dry, c. 14-20
mm. long, usually shortly apiculate, sometimes rounded or obtuse at the apex, roun-
ded to acute at the base, glabrous; endocarp very slightly longitudinally ridged,
very slightly flattened but without prominent lateral ridges.

VENEZUELA: Amazonas: Cerro Sipapo, ‘“‘ buttressed tree 30 m. high, 40 cm.
diam., dense low elevation forests ’’, 24 Nov. 1948, Maguire & Politi 27370 (BM).

BraziL: Amazonas: Estrado do Aleixo, Manaos, in non-flooded forest, 14 June
1933, Ducke 25016 (K). Cachoeira Grande, Manaos, in non-flooded forest, 3 Jan.
1937, Ducke 374 (K). Mandaos, Rio Taruma, “terra firme, alta, floresta alta’’,
10 Sept. 1950, Frdes 26580 (BM). Parad: Vila Braga, Rio Tapajoz, medium-sized
tree in non-flooded wood by waterfall, 23 Sept. 1922 and 24 May 1923, Ducke 17686
(K ; RB, lectotype, not seen). Bosque Rodrigues Alves, Belém, without date,
Guedes 233 (BM). Belém, without date, Guedes 239 (BM).

Ducke also recorded the species “inter flumina Cumina-Mirim et Ariramba affl.
Rio Trombetas ”’, Ducke 13584 (RB), and as sterile trees near Belém, Breves and
Faro, all in the state of Para.

But for Maguire G& Politi 27370, which was originally named B. capitata, this
species has been found only near the Amazon and its tributaries in northern Brazil.
The Venezuela specimen is from a locality close to the Rio Orinoco some 1,250 kilo-
metres from Manaos, the nearest Brazilian locality. The distinctive leaves, however,
make its identification quite definite. 7

No specimen we have seen has flowers, those of Ducke 17686 (which we have chosen
as lectotype) having just fallen. The ovary in this gathering is glabrous, and as
Ducke also states this it seems that this character provides a further good means of
distinguishing B. parvifolia from B. capitata. By far the greater number of Buchen-
avia species have densely pubescent ovaries, the indumentum often disappearing
to give quite glabrous fruits (e.g. B. oxycarpa). B. grandis appears to be the only
other species with a glabrous ovary.

3. Buchenavia kleinii Exell in Ann. & Mag. Nat. Hist., Ser. 12, 6 : 400 (1953).

Leaf-lamina chartaceous when young, subcoriaceous when mature, 2-9 X I°5-5
cm., obovate to broadly obovate, rounded or obtuse or subacute or apiculate at the
apex, acute-cuneate but not decurrent at the base, densely pubescent when very
young, becoming glabrous except on the midrib and lateral nerves when mature ;
major lateral nerves 4-8 pairs ; petiole distinct, fairly long in most cases, 8-20 mm.
long. Inflorescences as in B. capitata; peduncle rather stout, Io-20 mm. long.
Fruit ellipsoid or less often obovoid, 17-28 x 11-14 mm., apiculate or less often
rounded at the apex, densely but minutely rufous-puberulous, becoming less hairy
when very old ; endocarp very similar in shape to that of B. parvifolia.

BraZiIL: Santa Catarina: Mata do Hoffmann, Brusque, 13 Jan. 1951, Klein 22
(BM; S, holotype). Cunhas, Itajai, 10 m., 29 Nov. 1954, Klein 870 (BM) ; same

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 15

locality, 8 Feb. 1955, Klein 1144 (BM); same locality, 10 Mar. 1955, Klein 1182
(BM) ; same locality, 26 July 1955, Klein 1479 (BM). Bracgo Joaquim, Luis Alves,
Itajai, 300 m., 14 Dec. 1954, Klein 924 (BM) ; same locality, 350 m., 13 Jan. 1955,
Klein 1075 (BM). Horto Florestal I.N.P., Ibirama, 250 m., 27 Dec. 1954, Klein 940
(BM) ; same locality, 300 m., 4 Feb. 1956, Reitz & Klein 2568 (BM). Morro da
Fazenda, Itajai, 50 m., 17 Mar. 1955, Kleim 1232 (BM). Piloes, Palhoga, 300 m.,
1r Jan. 1957, Klein 2158 (BM). Mata da Companhia Hering, Bom Retiro, Blumenau,
250 m., 15 Dec. 1959, Klein 2342 (BM). Morro Spitzkopf, Blumenau, 700 m.,
1r Mar. 1960, Klein 2424 (BM). Morro da Ressacada, Itajai, 250 m., 29 Mar.
1956, Reitz & Klein 2915 (BM). Matador, Rio do Sul, 350 m., 27 Jan. 1959, Reitz &
Klein 8319 (BM). Alto Matador, Rio do Sul, 800 m., 30 Dec. 1958, Reitz 6126
(BM). Vargem Grande Lauro Miiller, 350 m., 20 Feb. 1959, Reitz & Klein 8494
(BM).

ae species is known only from the south-eastern corner of Santa Catarina,
southern Brazil, although it may well occur elsewhere. The area is south of the
Tropic of Capicorn, B. kleini: being the only species of the genus (and one of the
comparatively few members of the Combretaceae) known outside the tropics. Its
vernacular name is given as ‘‘ Guarajuba ”’.

B. kleinii appears to be most closely related to B. capitata, but its puberulous fruits
and characteristic leaf-shape make it quite easily identifiable. In the shape of its
endocarp it approaches B. parvifolia more closely than B. capitata.

4. Buchenavia sericocarpa Ducke in Bol. Técn. Inst. Agron. Norte 4 : 23 (1945).

Leaf-lamina subcoriaceous, 3-12 X I-6 cm., narrowly elliptic to elliptic, short-
to rather long-acuminate at the apex, cuneate and narrowed into the petiole at the
base, very sparsely pubescent to almost glabrous when mature (except for appressed
pubescence on the midrib and lateral nerves on both surfaces) ; midrib and lateral
nerves conspicuous and raised below; major lateral nerves 5-10 pairs; petiole
distinct, c. 7-30 mm. long, appressed-fuscous-pubescent. Peduncle fairly stout in
fruit, c. 20-25 mm. long, rather sparsely appressed-pubescent ; rhachis extremely
short, up to c. 8 mm. long. Fruit narrowly ellipsoid-oblong, more or less terete,
Cc. 20-25 X 7-I0 mm. at maturity, rounded to very broadly obtuse at both ends,
densely but shortly appressed-pale-tawny-pubescent.

BrAzIL: Amazonas: Manaos, Estrada do Bombeamento, ‘‘ Capoeirdo, terra
firme, arvore pequena ”’, 26 Nov. and 30 Dec. 1943, Ducke r48r (K ; RB, holotype,
not seen).

Known only from the type gatherings (presumably made from the same tree).

This species is very different from all others of the genus with a capitate inflores-
cence in its highly distinctive leaves. There is a remarkable similarity between this
species and B. acuminata, so much so that there is scarcely any other difference
that we can detect apart from the elongate inflorescences of the latter. This could
be a striking example of parallel evolution but would seem to be more probably

BOT; 35-3. 1§§§

16 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

the comparatively recent evolution of a species by the appearance of a single but
very distinctive character expressing one of the fundamental tendencies in the family
towards massing of the flowers in a condensed inflorescence. A close parallel is
shown in the relationship between Combretum punctatum subsp. sguamosum (Roxb.
ex G. Don) Exell and C. punctatum Blume subsp. punctatum, but in that case the
difference in the form of the inflorescence is not so clear-cut and is considered to
be of no more than subspecific value.

5. Buchenavia ochroprumna Eichl. in Flora 49:165 (1866); in Mart., FI.
Brasil. 14 (2) : 96 (1867).

Leaf-lamina subcoriaceous when mature, c. 2-9°5 X I-4°5 cm., oblanceolate to
obovate, usually rounded but sometimes very broadly obtuse or (when damaged?)
retuse at the apex, cuneate and somewhat decurrent into the petiole at the base,
almost glabrous when mature except on the midrib and lateral nerves; major
lateral nerves 3-7 pairs; petiole rather short, 4-15 mm. long, usually rather stout.
Peduncle 6-30 mm. long, slender at flowering time, stout in fruit. Fruit ellipsoid
or ovoid, indistinctly 5-angled, markedly and irregularly ridged when dry, 18-25
x 7-15 mm. including beak, long-beaked at the apex, rounded at the base, densely
but minutely ferrugineous-tomentose ; beak 4-8 mm. long, narrow, straight or
somewhat curved; endocarp markedly longitudinally ridged (more so than in
B. capitata), often about 5 ridges more prominent than the rest.

BRAZIL : Amazonas: Manaos, Igarapé da Cachoeira Grande, “igapd profunde
inundato, arbor parva’’, 28 Mar. 1937, Ducke 445 (K) ; 28 Mar. 1937, Ducke 35172
(K). Mandos, “‘igapo ad faucem fluminis Taruma Grande’’, 7 Apr. 1932, Ducke
25019 (K). Rio Negro, in the vicinity of Barcelos, 19 Apr. 1952, Frées 28438 (BM).
Region of Parintins, Lago do Juruti, 18 Jan. 1957, Frdes 33069 (BM). Parad:
Near Santarém, Jan. 1850, Spruce 309 (M, lectotype) and without n. (BM; K).
Santarém, by river, “‘ bush of ro feet’, Jan. 1850, Spruce 619 (K). Lago Cugari,
plateau of Santarém, 15 Apr. 1955, Frdes 31671 (BM). Igarapé Cugari, plateau of
Santarém, 15 Apr. 1955, Frdes 31773 (BM). Mata da Pirelli, Ananindeua, 8 Jan.
1958, Silva 501 (BM). Rio Aramun, Almeirim, 2 Sept. 1918, Ducke 17287 (BM).

This species is thus known only from the lower regions of the Amazon. Eichler
cited only Spruce 309, which he saw in Herb. Leningrad and Herb. Munich. We have
seen the latter specimen, which we select as the lectotype. Ducke 17287, labelled
B. oxycarpa, is almost certainly B. ochroprumna as shown by the rather distinctive
leaves. Besides their shape, the leaves of B. ochroprumna are usually characterized
by a ferrugineous colour on their under surfaces when dry. This specimen is the
only one we have seen in flower. The flowers do not appear to differ from those of
B. capitata.

B. ochroprumna differs from all other species of the genus in its highly character-
istic fruits, and may also be distinguished by its leaves.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 7

6. Buchenavia punctata Eichl. in Flora 49: 166 (1866); in Mart., Fl. Brasil.
14 (2) : 98 (1867).

Terminalia punctata Spruce ex Eichl., loc. cit. (1867), nom. syn.; non T. punctata Roth
(1821) nec Eichl., tom. cit. : 85 (1867).

Leaf-lamina chartaceous when mature, c. 6-I5 X 2:5-4:5 cm., conspicuously but
minutely pellucid-punctate, very narrowly obovate to oblanceolate, acute to sub-
acute at the apex, very narrowly cuneate but not decurrent into the petiole at the
base, very sparsely pubescent above, pubescent on the midrib and nerves and sparsely
pubescent elsewhere below ; midrib and major lateral nerves raised and very con-
spicuous below; reticulation rather inconspicuous ; major lateral nerves c. 7—II
pairs; petiole short and stout, 5-15 mm. long, eglandular, pubescent. Peduncle
stout, c. 35 mm. long; rhachis long (over 25 mm., but material fragmentary). Fruit
ellipsoid, radially symmetrical, markedly 5(6)-ridged along its length, 27-31 x 12-19
mm., conspicuously apiculate at the apex, narrowed into a very short but distinct
pseudostipe at the base, very sparsely pubescent between the ridges, glabrous else-
where ; apiculus 1-3 mm. long.

PERU: San Martin: In rocky stream, near Tarapoto, ‘“‘ Arbor 25 pedalis ramis
paucis longis validis brevi-ramulosis ’’, 1855-56, Spruce 4945 (BM; BR, lectotype,
not seen; K).

This species is very distinct from all others of the genus in both its fruits and
leaves. Only B. pterocarpa has fruits as conspicuously ridged. Since the flowers
are unknown we cannot, of course, be absolutely certain that this species belongs to
Buchenavia, but as the fruit is distinctly less winged than that of any American species
of Terminalia we have seen it seems safe to include it in Buchenavia. Its locality
is completely isolated from that of any other Buchenavia yet discovered and is one
of only two records of the genus for Peru.

7. Buchenavia reticulata Eichl. in Flora 49: 166 (1866); in Mart., Fl. Brasil.

14 (2) : 98 (1867).

Leaf-lamina more or less chartaceous when mature, (2) 5-26 (36) x I-7 (12) cm.,
narrowly elliptic to oblanceolate, obtuse to acute or shortly apiculate at the apex,
narrowly cuneate but not decurrent at the base, ferrugineous-tomentose over the
whole of the lower surface at flowering time and during fruit maturation but tomen-
tum wearing off after fruit-fall, ferrugineous- to rubiginous-tomentose or -pubescent
on midrib and lateral nerves of upper surface at flowering time but indumentum
wearing off during fruit maturation, sparsely pubescent and becoming glabrous
elsewhere on upper surface ; midrib very conspicuous and markedly raised below ;
major lateral nerves conspicuous and markedly raised below, 5-15 pairs, arising at
c. 45° or less ; reticulation at and after flowering time prominent and raised below,
mostly consisting of secondary venules connecting the major lateral nerves ; petiole
distinct but usually short, c. 6-23 mm. long, stout, ferrugineous- or rubiginous-
tomentose, most often conspicuously biglandular towards the apex. Peduncle

18 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

fairly stout, c. 20-30 (40) mm. long, ferrugineous- or rubiginous-tomentose ; rhachis
c. 50-140 mm. long, ferrugineous- or rubiginous-tomentose ;_ flowers rather large,
c. 4-5 mm. long. Lower hypanthium ferrugineous- to rubiginous-tomentose round
the ovary, almost glabrous and c. 1-2 mm. long above it ; upper hypanthium c.
I'5-2 X 2°5-3°5 mm., almost or quite glabrous outside. Fruit narrowly ellipsoid
to ellipsoid, irregularly and rather slightly longitudinally furrowed or scarcely so,
sometimes slightly flattened, c. 19-28 x 8-12 mm., apiculate to acuminate at the
apex, shortly (1-2 mm.) pseudostipitate at the base, shortly ferrugineous- to rubi-
ginous-tomentose and rather velvety all over ; apiculus 1-6 mm. long, straight or
slightly curved ; endocarp variable, very slightly to conspicuously longitudinally
furrowed, more or less circular in section to rather flattened, sometimes with 2 very
narrow lateral wings.

VENEZUELA: Amazonas: On the Rivers Casiquiare, Vasiva and Pacimoni,
1853-54, Spruce 3453 in part (BM ; BR, lectotype, not seen; K). Along the River
Pacimoni, frequent, and also frequent along the Rivers Guainia and Vasiva, Feb.
1854, Spruce 3453 in part (K, isolectoparatype). Frequent on banks of the River
Guainia, “‘ Arbor parva patula 18 pedalis. Fructus molles ovali-fusiformes acuti
haud angulati. This and Parkia auriculata are perhaps the commonest riparial
trees of the rivers Guainia, Pacimoni, Vasiva and Atabapo”’, May 1854, Spruce
3453 in part (K, isolectoparatype). Cafio Yapacana, Cerro Yapacana, Alto Rio
Orinoco, “‘ tree 3-10 m. high, fruit brown, frequent at laguna edge’, 19 Mar. 1953,
Maguire & Wurdack 34584 (BM). Frequent along Caio Yapacana, below port for
Cerro Yapacana, Rio Orinoco, “ tree to 20 m. high, inflorescence brown ’’, 6 Jan.
1951, Maguire, Cowan & Wurdack 30760 (BM).

BRAZIL: Amazonas: Above Kibaru, Rio Negro, 23 June 1874, Tvaill 216 (K).
Above Santa Izabel, Rio Negro, 5 Mar. 1936, Ducke 35167 (K). Riosinho Juruema,
Alto Rio Jutahy, Solimoes, “ Terra firme, igapés. Arvore’’, 4 June 1945, Foes
21039 (K). Igarapé Imuta, tributary of Rio Negro, ‘terra firme. Arvore de 8 m.,
ramos inclinados ; flores roseas’’, 18 Mar. 1952, F’vdes 27936 (BM). Rio Negro, May
1947, Froes 22323 (BM).

From Spruce’s records for Rio Guainia and Rio Atabapo it seems probable that
this species also occurs in Colombia (see footnote under Ramatuella argentea con-
cerning Spruce 3498, p. 41 below).

This very distinct species has very large or large leaves although it is a rather
small tree. It is apparently abundant in the upper Rio Negro-Orinoco region
(fide Spruce 3453), and thus entirely separated from the other two species with
reticulate tomentose leaves (B. tomentosa and B. callistachya) which occur only in
the lower regions of the Amazon basin and further south (Fig. 2).

The fruits of this species show a remarkable range in the form of the endocarp.
In some specimens it is round in section and only slightly longitudinally furrowed,
while in others it is conspicuously flattened, markedly longitudinally ridged, and has
two lateral wings, although these are not as well developed as in B. pterocarpa.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELEA 19

8. Buchenavia tomentosa Eichl. in Flora 49: 166 (1866); in Mart., Fl. Brasil.
14 (2) : 97 (1867).—Glaziou in Mém. Soc. Bot. France 1 (3) : 203 (1908).
Terminalia tomentosa Mart. ex Eichl., loc. cit. (1867), nom. syn.; non T. tomentosa

Wight & Arn. (1834).
Buchenavia corrugata Ducke in Arch. Jard. Bot. Rio Janeiro 4 : 150 (1925).

Leaf-lamina chartaceous at flowering time, coriaceous when mature, c. 3°5-
13°5 X I°5-7 cm., narrowly elJiptic or oblanceolate to (probably always at maturity)
elliptic or obovate, rounded at the apex but obtuse or more pointed when very
young, cuneate at the base, often inrolled at the margin, ferrugineous- or rubiginous-
tomentose or densely pubescent all over both surfaces at flowering time, indumentum
beginning to wear off soon afterwards ; midrib very conspicuous and markedly raised
below ; major lateral nerves conspicuous and somewhat raised below, c. 9-14 pairs,
arising at about 45°; reticulation not prominent before flowering time but rather
conspicuously so afterwards, though less so than in B. reticulata and B. callistachya,
less regular than in B. reticulata ; petiole distinct, c. 3-18 mm. long, stout, densely
rubiginous- or ferrugineous-velutinous when young, sometimes biglandular. Peduncle
c. 6-20 mm. long, usually stout, densely rubiginous-velutinous when young ;
rhachis c. 3-11 cm. long, densely rubiginous-velutinous when young ; flowers very
like those of B. reticulata, c. 4-5 mm. long, indumentum identical. Upper hy-
panthium c. 2-5-4 mm. across. Fruit ellipsoid to more or less spherical, probably
more or less terete when fresh, 22-31 X I5-25 mm., more or less rounded at both
ends, rather sparsely ferrugineous-pubescent when mature.

Brazi_: “ Brasilia’’, Pohl 83 (W, lectoparatypes). Pard: Serra Pontada, in
the Jutahy region between Almeirim and Prainha, “silva, arbor magna’’, 11 Sept.
1923, Ducke 17677 (K, isotype of B. corrugata). Santa Cruz dos Martirios, regiao do
Araguais, “‘ arvore de ro m., frutos comestiveis ’’, 15 June 1953, F'vdes 30018 (BM).
Piaut: Banks Gurgia (= Rio Gurgeia ?), “a large tree called Biriba—fruit acrid
and bitter—liked by deer ’’, Aug. 1839, Gardner 2657 (K). Goids : Between Alegre
and Ponte do Sevorino, “ arbre de petite taille ’’, 2 Oct. 1895, Glaziou 21125 (K; P).
In woods and plains, Serra da Chapada, “ arbor 15-20 ped.”’, Sept. 1827, Martius
1180 (LE, lectoparatype). Minas Gerais: Biribiry (fide Glaziou), 1892, Glaziou
19144 (K). On sandy plains between Alegres and Rio Sao Francisco, “‘ arbor 8-20
ped.”’, Sept. 1834, Riedel 2641 (BM; K; LE, lectotype and lectoparatype ; W).

Glaziou 21125 is unlocalized in Herb. Kew, but the Paris specimen bears a label
concurring with the locality cited by Glaziou (1908). This sheet also bears a label
identical to that of B. macahensis (nomen nudum ; see p. 37 below), Glaziou 18218,
but the actual specimen bears a tag numbered 21725 and other pieces of evidence
conclusively show that the presence of the second label is due to an error in mounting.

Eichler (1866) cited specimens of B. tomentosa in Herb. Munich, Herb. Martius,
Herb. Vienna and Herb. Leningrad, but those in the first two herbaria do not appear
to have survived. Of the three specimens in Herb. Vienna (W) one is a duplicate
of one of the Leningrad specimens whilst the other two (both Pohl 83) are mounted
on the same sheet. There are also three specimens in Leningrad : one collected by
Martius and two (duplicates) by Riedel. We have selected the better Riedel

20 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

specimen as the lectotype as duplicates of this are also present in the British
Museum, Kew and Vienna Herbaria.

B. corrugata shows considerable similarity to B. callistachya, but the differences
noted by Ducke and ourselves appear to be sufficient to keep the two separate, at
least for the present. Comparison of them with the type specimens of B. tomentosa
leads us to the opinion that B. corrugata is conspecific with the latter.

B. tomentosa seems to be a species which does not extend up the Amazon, in con-
trast with B. reticulata which is confined to the upper Amazon and Orinoco
tributaries (Fig. 2). The vernacular name is noted as ‘‘ Tanebuco ”’ by Pohl.

g. Buchenavia callistachya Ducke in Arch. Inst. Biol. Veg. Rio Janeiro 2 : 64
(1935).

Leaf-lamina coriaceous when mature, c. 5*5-I0°5 X 2°5-5°5 cm., elliptic to slightly
obovate, rounded at the apex, cuneate at the base, inrolled at the margin when
young, lower surface at flowering time densely ferrugineous-pubescent except tomen-
tose on the midrib and nerves, indumentum beginning to wear off soon afterwards
but midrib and nerves densely pubescent below even after fruit-fall ; midrib very
conspicuous and markedly raised below ; lateral nerves also markedly raised below,
c. 8-11 pairs, arising at 45° or slightly more ; reticulation very prominent below
after flowering time, less regular than in B. reticulata ; petiole distinct, stout, ferru-
gineous- to rubiginous-tomentose at flowering time, c. 6-16 mm. long, sometimes bi-
glandular. Peduncle c. 10 mm. long or less at flowering time, up to 15 mm. long in
fruit, stout, densely rubiginous-velutinous at flowering time, densely pubescent in
fruit ; rhachis 15-25 mm. long at flowering time, up to 45 mm. in fruit ; flowers
smaller than in B. reticulata and B. tomentosa, c. 2-5-3 mm. long, indumentum as in
the two preceding species. Upper hypanthium c. 1-75-3 mm. across. Anthers
also smaller. Fruit ellipsoid wrinkled when dry, probably more or less terete when
fresh, 21-23 X I2-I5 mm., rounded at the apex, rounded or very shortly pseudo-
stipitate at the base, finely ferrugineous- to rubiginous-puberulous when mature.

BraziL: Amazonas: Manaos, near Estrada do Aleixo, in non-flooded wood,
‘“ Arbor sat. magna, spicae anthesi pallide viridiflorae pedunculo brunneo-purpureo”’,
g July and 26 Nov. 1932, Ducke 25021 (K; RB, holotype, not seen). Maranhao :
Ilha dos Botes, Rio Tocantins, near Carolina, 25 May 1950, Pires & Black 2095
(BM).

B. callistachya is clearly closely allied to B. tomentosa, from which it differs in a
few characters which remain constant with regard to the rather few specimens of
each species known. The recently collected fruiting specimen (Pires & Black 2095)
almost certainly belongs to this species, its leaves being very similar to those of
the type. It is just possible, however, that we have confused the fruiting specimens
of B. callistachya and B. tomentosa, or that B. callistachya itself may prove to be
conspecific with the latter. These points must await more and better material for
elucidation.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 21

10. Buchenavia suaveolens Eichl. in Flora 49 : 166 (1866) ; in Mart., Fl. Brasil.
14 (2) : 97 (1867). (Fig. 3 e, f)
Buchenavia oxycarpa Eichl. in Flora 49: 165 (1866) pro parte, quoad Schomburgk 854 ;
in Mart., loc. cit. (1867) pro parte, quoad Schomburgk 854.
Terminalia suaveolens Spruce ex Eichl., loc. cit. (1867), nom. syn.

Terminalia vasivae Spruce ex Eichl., loc. cit. (1867), nom. syn. (Spruce also used a third
manuscript name which was not cited by Eichler.)

Leaf-lamina coriaceous when mature, 1-8-8 x 0-3-4:3 cm., narrowly elliptic or
oblanceolate to elliptic or obovate, retuse to rounded at the apex, narrowly to very
narrowly cuneate and somewhat decurrent into the petiole at the base, almost
glabrous when mature except on the midrib; midrib fairly conspicuous, slightly
raised below ; major lateral nerves inconspicuous to very inconspicuous, but usually
over 6 pairs visible below, scarcely or not raised ; reticulation not visible ; petiole
fairly long and distinct, c. 3-20 mm. long, very sparsely and minutely pubescent,
eglandular. Peduncle usually rather slender, sparsely pubescent, up to c. 25 mm.
long ; rhachis up to 65 mm. long at flowering time, sparsely pubescent; flowers
scarcely different from those of B. capitata, but ovary usually rather more sparsely
pubescent. Fruit ellipsoid to slightly obovoid, more or less terete when fresh,
24-30 X IO-I4 mm., conspicuously apiculate at the apex, narrowed into a short
but distinct pseudostipe at the base, glabrous, the apiculus 1-3 mm. long, more or
less straight ; endocarp ellipsoid, almost round in section, c. 20 X 9 mm., obtuse
at each end, more or less smooth.

VENEZUELA: Amazonas: On the rivers Casiquiare, Vasiva and Pacimoni, 1853-
54, Spruce 3198 (BM; K; LE, lectoparatype). On the banks of Lake Vasiva,
also observed on Rio Pacimoni, Dec. 1853, Spruce 3190 (K). Rio Sanariapo, 120 m.,
2 July 1942, Willams 15955 (F).

VENEZUELA or COLOMBIA: On the banks of Rio Guainia, “‘ Drupa viridis carnosa,
recens vix obsolete prismatica’”’, June 1854, Spruce 1887 in part (K). On Rio
Guainia above its confluence with Rio Casiquiare, 1854, Spruce 1887 in part (K).
(See footnote under Ramatuella argentea concerning Spruce 3498, p. 41 below.)

BrAziL: Amazonas: Rio Negro, between Barra and Barcellos, Nov. 1851,
Spruce 1887 in part (BM; K; LE, lectotype). Caloruca, Rio Preto, tributary
of Rio Negro, 19 Apr. 1952, Frdes 28309 (BM). Rio Branco: ‘‘ Guiana ”’ (but actu-
ally by Falls on the Rio Branco, fide Schomburgk, MS.), 1839, Schomburgk 854
(BM; K). “Guiana Britannica’’, 1839, Schomburgk 814 (BM) (probably error
for 854).

B. suaveolens is thus confined, like B. pterocarpa and B. reticulata, to the upper
Amazon-Orinoco region (Fig. 2).

The distinction between B. suaveolens and B. plerocarpa is made under the latter
species.

Eichler (1866) cited specimens in Herb. Leningrad and Herb. Martius, of which
we have seen the former. The two gatherings are Spruce 1887 and Spruce 3198
(which Eichler gives in error as 3789), and there are duplicates of both at the
British Museum and Kew. We have selected Spruce 1887 as the lectotype as there

22 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

LMR "4

Fic. 3. Buchenavia pterocarpa Exell & Stace (holotype) : a, habit (x %) ; 6, c, two views
of fruit (x1); d, fruit in transverse section (x1). B. suaveolens Eichl. (Frdes 28309) :
e, fruit (x1); f, fruit in transverse section (x 1).

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 23

is no confusion about the number and other specimens (from another locality) also
with this number bear fruits.

Schomburgk 854 is labelled ‘“‘ Guiana’’ but according to the collector’s note-
books he was actually at the Falls of the Rio Branco, Brazil. Schomburgk 814 is
similarly labelled and since his note-book states that this number was a herbaceous
plant it seems most probable that 8r4 is an error for 854, which it resembles in every
detail. B. suaveolens is otherwise not known from British Guiana. Eichler (1866,
1867) cites Schomburgk 854 under B. oxycarpa (in 1867 as an unnamed variety),
but this is clearly a misidentification.

11. Buchenavia pterocarpa Exell & Stace, sp. nov. (Fig. 3 a-d.)

Arbor parva ad c. Io m. alta; ramuli griseo-fusci, fere glabri, apice non clavati.
Folia spiraliter ordinata ad ramulorum apices congesta; lamina subcoriacea,
I*5-9°5 X 0°5-2°8 cm., anguste elliptica vel oblanceolata ad anguste obovata, in-
tegra, apice obtusa rotundata vel retusa, basi acuta decurrens, fere glabra, costa
media infra prominenti, nervis lateralibus inconspicuis; petiolus c. 3-18 mm.
longus, sparse minuteque puberulus eglandulosus. IJnflorescentiae axillares, elongato-
spicatae ; pedunculus gracilis, c. to-17 mm. longus, sparse minuteque puberulus ;
rhachis c. 15-53 mm. longa, puberula ; flores ferrugineo-virides ; bracteae parvae,
ante anthesin caducae. Hypanthium inferum gracile, I-2 mm. longum, ovario in
parte basali puberula incluso, parte apicali sparse minuteque puberula ; superum
Cc. I-I'5 X 2-3 mm., extus sparse puberulum, intus pubescens, margine saepe fere
integrum. Stamina 10, ad o-5-r mm. exserta. Stylus ad c. I°5 mm. exsertus,
filiformis, glaber. Fructus (plerumque singulus tantum pro infructescentia matur-
escens) latissime applanato-ellipsoideus vel obovoideus, c. 14-28 * 10-16 mm. et 5-8
mm. crassus, apice abrupte apiculatus, basi rotundatus vel breviter (ad 3 mm.)
pseudostipitatus, glaber, apiculo ad 4 mm. longo curvato, alis 2 lateralibus angustis
et angulis 3 prominentibus instructus ; endocarpium lignosum, lateraliter bialatum,
irregulariter longitudinaliter sulcatum.

VENEZUELA: Amazonas: Occasional along Rio Yatua, near Laja Catipan,
Casiquiare, 100-140 m., “ flat-topped riverine tree, 6-8 m. high, fruit green ’”’,
6 Feb. 1954, Maguire, Wurdack & Bunting 37543 (BM, holotype). Common and
subdominant along Cana Catua, Cerro Yapacana, Rio Orinoco, 125 m., 19 Nov.
1953, Maguire, Wurdack & Bunting 36555 (BM). Cano Yapacana, below port to
Cerro Yapacana, Rio Orinoco, 6 Jan. 1951, Maguire, Cowan & Wurdack 30763
(BM).

BRAZIL: Amazonas : Santa Izabel, Rio Negro, in flooded riparian forest, 8 Oct.
1932, Ducke 25018 in part (K) ; same locality, 9 Mar. 1936, Ducke 25018 in part
(K). Igarapé Imuta, tributary of Rio Negro, ‘‘ terra firme’”’, 12 Mar. 1952, Froes
27937 (BM).

All the specimens we cite, other than Frydes 27937, were previously labelled
B. suaveolens. In the absence of fruits it does not always seem to be possible to

24 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

distinguish between B. suaveolens and B. pterocarpa, but in most cases the leaves
of the former are broader and more markedly obovate. The fruits of the two
species are, however, quite different (see Fig. 3). The fruit of B. suaveolens is more
or less terete, narrower in proportion to its length and with a usually straight apiculus
abruptly delimited from the obtuse or rounded apex. In our new species on the
other hand the fruit is considerably flattened with two narrow lateral wings and three
extra ridges (one on one face and two on the other), is broader in comparison with
its length and has a usually curved apiculus very abruptly delimited from the
usually rounded or truncate or even retuse apex. The endocarp is also markedly
different in the two fruits, that of B. plerocarpa having two lateral wings and an
irregularly furrowed surface, while that of B. suaveolens is smooth and almost round
in cross-section. The bilateral symmetry of the fruit of B. pterocarpa is not found
so greatly developed elsewhere in the genus, although some species (notably B. reti-
culata and less conspicuously B. parvifolia and others) show tendencies towards it.

Comparison of recent collections of B. suaveolens and B. pterocarpa with Spruce’s
collections of the former shows that it is the flattened-fruited species which needs
to be described as new. Although Eichler did not see fruits of B. suaveolens (none
of his cited specimens is in fruit, and he states “‘ Fructus ignotus ”’ in Fl. Brasil.) it
seems clear that it was the terete-fruited species that he described. The evidence
for this is that both Leningrad specimens are broad-leaved, and that other specimens
of Spruce 1887 (not duplicates of those which Eichler cited from Barra and Bar-
cellos) from the Rio Guainia which are in Herb. Kew possess the distinctive terete
fruits.

12. Buchenavia oxycarpa (Mart.) Eichl. in Flora 49 : 164, t. 3 figs. 17, 18, 20, 21
(r866) excl. Schomburgk 854; in Mart., Fl. Brasil. 14 (2) : 97, t. 25 (1867) excl.
Schomburgk 854.

Terminalia oxycarpa Mart. in Flora 24, Beibl. 2 : 22 (1841).

Leaf-lamina chartaceous, I-13 X 0°5—5 cm., very narrowly elliptic or oblanceolate
to obovate, rounded to acute at the apex, narrowly cuneate but scarcely decurrent
at the base, almost glabrous when mature except on the midrib ; midrib fairly con-
spicuous, slightly raised below ; major lateral nerves fairly conspicuous, very slightly
raised below, (4) 6-9 (12) pairs ; reticulation fine but fairly conspicuous ; petiole very
distinct, 4-25 mm. long, sparsely and minutely pubescent to almost glabrous, eglan-
dular. Peduncle slender, rather sparsely pubescent, up to 55 mm. long; flowers
very similar to those of B. capitata but ovary pubescent or more usually very sparsely
pubescent to almost glabrous, and always glabrous soon after flowering. Fruit
ellipsoid, wrinkled and slightly flattened to conspicuously 5-6 (7)-angled when dry,
19-30 X 9-I5 mm., abruptly acuminate at the apex, rounded to obtuse at the base,
glabrous ; acumen 3-9 mm. long, pointed, more or less straight or slightly curved ;
endocarp strongly longitudinally ridged, 5 ridges stronger than the rest, more or less
radially symmetrical but 2 lateral ridges sometimes the most prominent.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA_ 25

Brazi_: “Brasilia’’, without date, Martius (M, lectoparatype). ‘‘ Brazil’’, with-
out date, Poeppig 2689 ex Herb. Van Heurck (BM). “‘ Brasilia’’, without date,
Riedel 744 ex Herb. Hort. Petrop. (K). Amazonas: In inundations on the bank of
the river Solimoes, near Ega and elsewhere in the province of Rio Negro, 1820, Martius
(M, lectotype). ‘‘ Prov. Rio Negro prope Ega ad ripam fluv. Solimoes ’’, without
date, Martius (K; M; lectoparatypes). ‘In ripa fluv. Solimdes haud infrequens’’,
1820, Martius (M, lectoparatype). Humayta, near Tres Casas, basin of Rio Madeira,
“tree 40 ft. high’’, 14 Sept.—rz Oct. 1934, Krukoff 6174 (BM; K). Humayta,
near Livramento, on Rio Livramento, basin of Rio Madeira, on immediate shore
of river, “shrub 20 ft. high’, 12 Oct.—6 Nov. 1934, Krukoff 6733 (BM; F; K).
Parana do Careiro (mouth of the Solimées), “‘ Arvore pequena ; flér pardo vermelho
com estamos brancos’’, 26 Oct. 1946, Ducke 2023 (BM). S. Paulo de Olivenga,
igapé de Camatia, 2 Feb. 1937, Ducke go2 in part (K). Para: Near Santarém
and Paricatuba, “silvae ad fluv. Amazonum locis inundatis’’, without date,
Martius 2852 in part (M, lectoparatype). Near Santarém and Paricatuba, “In
sylvis ad fluv. Amazonum in locis inundatis”’, without date, Martius (M, lecto-
paratype). Near Santarém, Nov. 1849—Mar. 1850, Spruce 127 (M) and without n. (BM ;
K). Rio Tapajoz, Santarém, “‘ small tree, lvs. and fruits fastigiate on short ramuli”’,
Mar. 1850, Spruce 729 (K). On varzia land near Cassipa, Tapajoz River region,
Sept. 1931, Krukoff 1241 (BM; K). Pard/Minas Gerais : “ Para et Minas Geraes,
in sylvis ad rip. Amazonum, nec non Rio Fermozo’’, without date, Martius 2852
in part (mixed with Combretum laurifoliwm Mart.) (M, lectoparatype).

Peru: Loreto: Inundation belt of Rio Itaya, Iquitos, “‘ Shrub, flowers reddish-
brown ”’, 4 Nov. 1946, Asplund 14302 (K).

Out of Martius’s eight syntypes we have selected a specimen possessing leaves,
flowers and fruits as the lectotype.

B. oxycarpa is a very distinct species, although Eichler (1866, 1867) cited a gather-
ing of B. suaveolens (Schomburgk 854) under it. The endocarp of the present species
is markedly ridged and more or less radially symmetrical, a feature only as greatly
developed elsewhere in B. punctata. The fruits are markedly ridged in some speci-
mens (e.g. in Martius’s syntypes) and in Eichler’s figures. Spruce’s specimens
show no sign of these ridges, having slightly flattened and minutely wrinkled fruits.
Fruits of Ducke 402 are, however, more or less intermediate in this feature. Again,
in Krukoff 1241 and Krukoff 6174 the ovaries are completely glabrous at anthesis
or just after, while those of Asplund 14302 are ferrugineous-tomentose. The ovaries
of the flowers of Riedel 744 are completely intermediate in pubescence. The in-
florescences of Asflund 14302 are additionally unusual in that the whole peduncle
and rhachis are tomentose ; the long pedicel-like distal portion of the lower hypan-
thium and the outside of the upper hypanthium are, however, glabrous.

Eichler also recorded the species from the state of Mato Grosso (on banks of Rio
Paraguay), Brazil. Since he cites Riedel as the collector from this locality, the
specimen in question may well be the Riedel specimen referred to above under un-
localized records for Brazil.

26 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

13. Buchenavia discolor Diels in Verh. Bot. Ver. Prov. Brandenb. 48: 192 (1907).

Leaf-lamina chartaceous or coriaceous, c. 6-7-5 x 4.cm., obovate, obtuse to emar-
ginate at the apex, narrowly cuneate at the base, sparsely pubescent below ; major
lateral nerves prominulous below, 3-4 pairs, venules inconspicuous; petiole 10-15
mm. long. Peduncle plus rhachis 20-30 mm. long. Fruit ellipsoid, 5-angled, c.
15 mm. long excluding acumen, abruptly acuminate at the apex, tomentellous ;
acumen up to 10 mm. long. (Ex descr. orig.)

BRAZIL: Amazonas: Near Mandaos, on the banks of Rio Negro, Dec. 1901, Ule
5979 (B, holotype, destroyed).

The only known specimen, which was in the Berlin Herbarium, has been destroyed.
The species appears to be very distinct. Diels states that B. discolor is close to
B. oxycarpa but differs in the structure of the leaves (mainly in having only three or
our pairs of lateral nerves) and in the tomentellous fruit. It is quite possible, if the
inflorescences have been misinterpreted by Diels, that this species is the same as
B. ochroprumna, which has capitate inflorescences.

14. Buchenavia megalophylla Van Heurck & Muell. Arg. in Van Heurck, Obs.
Bot. > 2r1 (1870).

Leaf-lamina subcoriaceous when mature, I4-3I X 5-10 cm., narrowly obovate
to oblanceolate, obtusely apiculate to shortly acuminate at the apex, very narrowly
cuneate at the base but not decurrent into the petiole, almost glabrous above except
on the midrib and lateral nerves, shortly pubescent on the midrib and lateral nerves
below, almost glabrous to very sparsely puberulous elsewhere below ; midrib very
prominent and raised below, slightly sunken above; lateral nerves conspicuous
below, (9) 10-14 pairs ; reticulation quite conspicuous, mostly connecting the lateral
nerves ; petiole distinct and stout, 12-23 mm. long, usually conspicuously bi-
glandular towards the apex, shortly pubescent. Peduncle rather stout, densely
puberulous, c. 20-30 mm. long ; rhachis fairly stout, densely puberulous or tomen-
tellous, c. 70-125 mm. long ; flowers rather large,c.4 x 3mm. Ovary tomentellous.
Upper hypanthium almost glabrous outside. Fruit very large, 30-48 mm. long,
rather abruptly narrowed into a long acumen at the apex and into a pseudostipe
at the base, ferrugineous-tomentellous ; body of fruit ellipsoid, 18-26 * II-I13 mm.,
slightly compressed, conspicuously 5-ridged ; pseudostipe often relatively long and
slender, 2~7 mm. long ; acumen rather various, straight or markedly curved, slender
or rather stout (5, damaged ?) ro-20 mm. long.

BRITISH GUIANA: Demerara: ‘‘ Demerara’’, without date, Parker (K, as Pamea
guianensis). Essequibo: Mazaruni Station, “shrub or small tree 8 ft. high from
sand bank by river ’’, rr Sept. 1942, Field No. F845 in part = Forest Dept. 3581 in part
(K) ; same locality, rr Apr. 1943, Field No. F845 in part = Forest Dept. 3581 in part
(K).

The type of this species is said to be in “ herb. van Heurck ”’ (AWH) but we have
been informed that it is not to be found there now. It was said (Van Heurck &

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 27

Muell. Arg., op. cit. : 212) to have been sent to Mlle. Hermine Reichenbach, probably
from Demerara, under the name Pamea guianensis Aubl., but collector and date are
not given. A specimen at Kew (cited above) bears the annotation ‘“‘ Mr. Parker
Demerara. Allosohabbo yielding excellent timber. Pamea guianensis. Aubl.
Terminalia Pamea. Aubl.” This was in Herb. Hooker, presented to Kew in 1867,
three years before Van Heurck and Mueller’s publication. It seems very likely
that the Kew specimen, with which Forest Dept. 3581 is obviously conspecific, is a
duplicate of the specimen described by Van Heurck and Mueller. However, in the
absence of confirmation of this and of the fact that Van Heurck’s specimen is actually
missing we cannot give the Kew specimen the status of a type.

A further difficulty is that in their lengthy description Van Heurck and Mueller
state that the inflorescences of this species are capitate, whilst the specimens we have
seen all have very long spicate inflorescences. The rest of the description, notably
of the leaves and fruits, fits the Kew Demerara specimen exactly. Without seeing
Van Heurck’s specimen it is not possible to account for this discrepancy with cer-
tainty, but it seems probably that the specimen described had incomplete inflor-
escences which had been broken off. We believe, like Sandwith (in sched.), that the
inflorescences of this species were misinterpreted by Van Heurck and Mueller, and
that they are in fact elongate spikes as stated in our description. There remains the
unlikely possibility, however, that B. megalophylla has a truly capitate inflorescence
and that the three Kew specimens represent a new species.

15. Buchenavia pallidovirens Cuatrec. in Fieldiana, Bot. 27 (1) : 107 (1950).

Leaf-lamina subcoriaceous, 3-II xX I-3-7 cm., oblanceolate to narrowly elliptic
or oblong-lanceolate, abruptly and sharply apiculate or mucronate or shortly
acuminate at the apex, narrowly cuneate but scarcely decurrent into the petiole
at the base, almost glabrous above except sparsely puberulous on the midrib and
lateral nerves, very sparsely puberulous or almost glabrous below except sparsely
pubescent or pubescent on the midrib and lateral nerves ; midrib conspicuous and
raised below ; lateral nerves conspicuous and raised below, 5-7 pairs, very dis-
tinctive, arising at about 40° or less to the midrib and then curving rather abruptly
inwards towards the apex and running at about 25° or less to the midrib for most of
their length and nearly parallel with the leaf margin at their apex ; reticulation con-
spicuous, very slightly raised below, forming very small inter-reticular spaces ;
petiole often long, 5-13 mm. long in flowering material, pubescent, obscurely bi-
glandular at the extreme apex in most leaves, 10-26 mm. long in fruiting material,
sparsely or very sparsely pubescent, not glandular or rather conspicuously bi-
glandular at about the middle. Peduncle 25-33 mm. long in flowering material,
8-18 mm. long in fruiting material, slender, sparsely pubescent, rhachis c. 40-70
mm. long, slender, pubescent ; flowers c. 3 X 2:5 mm. Ovary densely pubescent.
Upper hypanthium pubescent on the outside. Fruit (immature) ellipsoid-obovoid,
probably terete when fresh, up to 12 * 6 mm., subacute to obtuse at both ends,
glabrous.

28 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

CotomBIA: Valle: Costa del Pacifico, Bahia de Buenaventura, Quebrada de
San Joaquin, 0-10 m., “‘ Arbol grande. Racimos péndulos verdoso amarillentos.
Perianto verde amarillento claro. Anteras amarillas’’, 23 Feb. 1946, Cuatrecasas
19939 (BM; F, holotype, not seen).

BraziL: Amazonas: Rio Vaupés, Panuré, catinga, “‘ arbusto”’, 15 Nov. 1947,
Pires 1043 (BM).

In the above description ‘‘ flowering material’ refers to Cuatrecasas 19939 and
“ fruiting material ’’ to Pires 1043. These are the only two specimens known to us,
and it is of interest to note that they are from rather widely separated areas. It is
unfortunate that we have to compare a flowering specimen with a fruiting specimen
from a distant locality, but Pives 1043 seems most probably to belong to the same
species as the type. The peculiar nervation and reticulation of the leaves are com-
mon to both specimens and we have not met with them elsewhere in the genus.
The main point of difference is that the glands of Pives 1043 are, when present,
near the middle of the long petiole, while in Cuatrecasas 19939 they are either absent
or very inconspicuous and at the extreme apex of the petiole.

16. Buchenavia viridiflora Ducke in Arch. Inst. Biol. Veg. Rio Janeiro 2: 63
(1935).

Leaf-lamina subcoriaceous, I-I0 X 0°5-5:5 cm. (but rarely over 8 x 4 cm.).
oblanceolate or narrowly elliptic to obovate or elliptic, acute or subacute to rounded
or retuse at the apex, narrowly cuneate and often somewhat decurrent into the
petiole at the base, almost glabrous except sparsely to very sparsely pubescent on the
midrib and sometimes on the lateral nerves above, almost glabrous to sparsely
puberulous on the midrib and lateral nerves below ; midrib conspicuous and raised
below ; lateral nerves conspicuous and raised below, 4—7 pairs ; reticulation rather
inconspicuous ; petiole (4) 8-16 mm. long, pubescent, usually not glandular but
some very small (not young) leaves markedly biglandular at the apex of the petiole.
Peduncle 5-27 mm. long, slender, pubescent ; rhachis 12-50 mm. long, slender,
pubescent ; flowers 3-4:5 X 2-3 mm. Lower hypanthium densely pubescent
round the ovary, sparsely so above ; upper hypanthium almost glabrous outside.
Fruit more or less terete, up to 24 X I3 mm., subacute to very shortly apiculate at
the apex, rounded to subacute at the base, densely greyish-brown-tomentellous ;
endocarp terete, slightly longitudinally ridged, acute at the apex, rounded at the base.

BRAZIL: Amazonas: Near Cachoeira Grande, Manaos, forest in a high locality,
“arbor mediocris flor. laete viridibus ’’, 31 July 1932 and 8 Jan. 1933, Ducke 25022
(K). Estrada do Aleixo, Manaos, in non-flooded forest, 15 July 1932, Ducke 25023
(K ; RB, lectotype, not seen). Estrada do Aleixo, Manaos, in non-flooded forest,
26 Dec. 1936 and 19 Mar. 1937, Ducke 426 (K). Rio Branco: Boa Vista, Rio
Branco, non-flooded wood by Igarapé Caxanga, “ arbor mediocris floribus viridibus ”’,
24 Aug. (or July?) and 16 Sept. 1943, Ducke 1378 (F; K). Riverbank wood near
Boa Vista, Rio Branco, ‘ Bl. rétlichgelb, Bm. 5-15 m.”’, Oct. 1908, Ule 7682 (K).
Amapa: Rio Diapogue, Cachoeira Grande Rocha, “terra firme”, 19 Oct. 1950,
Froes 22698 (BM).

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 29

The leaf-shape of this species is apparently variable. In Ducke 1378, 25022 and
25023 the lamina is usually oblanceolate with a rounded apex, but some leaves on
the first of these have acute or subacute apices, showing a connexion with Ule 7682
which has larger, more or less elliptic acute leaves. Leaves on the fruiting branch of
Ducke 25022 (fruits are actually lacking in the specimen at Kew) are obovate and
rounded or retuse at the apex, which is the shape found in Frdes 22698. It seems,
however, that all the specimens cited belong to this species. Some of the inflor-
escences of the last specimen are very short, barely exceeding 3 cm. long with a
floriferous part little over r cm. long.

17. Buchenavia acuminata Exell & Stace, sp. nov.

Arbor ad 20 m. alta; ramuli griseo-fusci, sparse pubescentes, apice non clavati.
Folia spiraliter ordinata ad ramulorum apices congesta ; lamina chartacea, 2-9'5
x 0-8-5 cm., anguste elliptica vel elliptica, integra, apice obtuse vel acute acuminata
(acumine ad 15 mm. longo), basi acute cuneata, fere glabra costa media nervisque
lateralibus pubescentibus exceptis, costa media infra prominenti, nervis laterali-
bus infra prominentibus 5—8-paribus, venulis inconspicuis ; petiolus c. (4) 8-24 mm.
longus, pubescens, plerumque apice conspicue biglandulosus. IJmnflorescentiae
axillares, elongato-spicatae ; pedunculus gracilis, c. 15-30 mm. longus, sparse
pubescens ; rhachis gracilis, c. I5-30 mm. longa, sparse pubescens ; flores parvi ;
bracteae parvae, ante anthesin caducae. Hypanthium inferum gracile, c. 2 mm.
longum, ovario in parte basali dense pubescenti incluso, parte apicali pubescenti ;
superum Cc. I-I°5 X 2-2-5 mm., extus sparse puberulum, intus pubescens, margine
saepe fere integrum. Stamina Io, ad 0-75 mm. exserta. Stylus inclusus (semper?).
Fructus (pauci pro infructescentia maturescentes) ellipsoideus vel obovoideus, tere-
tiusculus, immaturus ad II x 5 mm., apice rotundatus, basi breviter pseudostipi-
tatus, dense argenteo-brunneo-tomentellus.

_ Brazit: Amazonas: Humayta, near Livramento, on Rio Livramento, basin of

Rio Madeira, on terra firma, “ tree 60 ft. high ’’, 12 Oct. -6 Nov. 1934, Krukoff 6916
(BM, holotype; F; K). Rio Urubu, Barreirinha, “terra firme’, ‘ Arvore de
7 m., frutos em espiguetas, amareladas ’’, 24 Aug. 1949, Frdes 25134 (BM).

The rather small long-acuminate leaves, long slender petioles and densely
tomentellous fruits make this species quite distinct from all others except B. serico-
carpa, which has a capitate inflorescence. Apart from this single but very con-
spicuous and easily recognizable difference we have been unable to separate these
two species, and they occur in the same area of Brazil. It is possible that here is
a case of a dimorphic species but until this is actually demonstrated it seems more
advisable to treat the plant with the elongate inflorescence as a different (new)
species.

18. Buchenavia fanshawei Exell & Maguire in Bull. Torr. Bot. Club 75 : 648
(1948).

Leaf-lamina coriaceous at maturity, I-14 x 0-5-6 cm., narrowly elliptic or ob-

lanceolate to elliptic or obovate, rounded to apiculate or less often acuminate or

30 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

acute at the apex, narrowly cuneate and often rather decurrent into the petiole at
the base, when mature almost glabrous above except sparsely pubescent on the
midrib, almost glabrous to very sparsely pubescent below except pubescent to
sparsely pubescent on the midrib and lateral nerves ; midrib conspicuous and raised
below ; lateral nerves conspicuous and raised below, 3-5 pairs ; reticulation rather —
inconspicuous or fairly conspicuous, not close; petiole slender to rather stout,
6-24 mm. long, pubescent to sparsely pubescent, usually not glandular but very
small leaves often conspicuously biglandular near top of petiole and some specimens
with normal leaves also biglandular ; some or most leaves on mature-leaved speci-
mens with conspicuous domatia (in the axils of the lateral nerves on the lower
surface) with a conspicuous opening smaller than the diameter of the domatium itself.
Peduncle slender at first, stouter in fruit, 6-30 mm. long, pubescent or puberulous ;
rhachis 7-38 mm. long, puberulous, often densely so ; flowers yellowish-green, c.
2°5-4 X 2-4 mm., seen on a rhachis up to only 20 mm. long. Fruit rather variable,
ovoid to ellipsoid- or oblong-ovoid but when ripe often becoming pressed almost
circular in outline, more or less terete when fresh or slightly flattened, up to 24 x 14
mm., subacute to rounded at the apex or rarely apiculate or shortly acuminate at
the apex with a beak up to 2 mm. long, rounded or shortly pseudostipitate at the
base, very shortly but rather densely tomentellous, indumentum wearing off when
very old; endocarp slightly flattened, slightly longitudinally ridged, acute to sub-
acute at both ends. _

‘

BRITISH GUIANA: Essequibo: Mazaruni Station, “ roo ft. tree 16 in. diam. from
mixed forest .. . fr. oval to oblong, yellow when ripe—pulp whitish, slightly sweet—
seed oval, whitish, veined, stone very hard’, 7 Sept. 1942, Field No. F844 = Forest
Dept. 3580 (K). Mazaruni Station, “ 60 ft. tree 10 in. diam. from secondary mixed
forest on brown sand... fls. in axill. revolute spikes, yellow-green—perianth shaped
like a flat bowl, barely lobed ’’, 5 May 1943, Feld No. F1270 = Forest Dept. 4006
(K). Mazaruni Station, seedlings of Forest Dept. 3580 ten weeks old from forest
nursery, no date, Field No. F'1616 = Forest Dept. 4352 (K). Mazaruni Station,
seedlings (no source) three weeks and seven months old from forest nursery, no date,
Field No. F2233 = Forest Dept. 4969 (K). Mazaruni Station, towards Labbakabra
Creek, 27 Aug. 1937, Sandwith 1219 (BM; K). Bank of Potaro River, Tumatumari,
4-6 July 1921, Gleason gor (K). Riverside below Tukeit, Potaro River Gorge,
rare, 16 May 1944, Maguire & Fanshawe 23499 (BM; K; NY, holotype). Mahdia
River, Potaro River, 172 km. along Bartica-Potaro road, 16 Jan. 1943, Field No.
F'1039 = Forest Dept. 3775 (K); same locality, 2r Jan. 1943, Field No. F 1064
= Forest Dept. 3800 (K). Basin of Kuyuwini River, tributary of Essequibo River,
about 240 km. from mouth, 12 Feb. 1938, Smith 3033 (K). Berbice: New River,
Courantyne River, 5 Oct. rg11, Anderson 749 (K).

BRAZIL: Amazonas: Cachoeira Baixa, Rio Taruma, Mandaos, “ 4arvore mediana ;
flr pardo avermelhado clara; fruto amarelado palido’’, May 1950, Ducke 2278
(BM).

Although not recorded from the third British Guianan county, Demerara, or from
Surinam, B. fanshawei probably occurs in both.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 31

B. fanshawet is a variable species, but is usually quite easily recognized in all
of its states. It appears to be most closely related to B. macrophylla, although in
their typical forms the two are extremely different in aspect. Some specimens of
B. macrophylla, however, have very small leaves with few nerves and stout inflores-
cences, while some specimens of B. fanshawei have acuminate leaves, biglandular
petioles and undeveloped or no domatia. The two species apparently overlap only
in the Mandos region of the middle Amazon (Fig. 2).

Leaves on more juvenile branches of B. fanshawei (but not juvenile leaves on older
branches) show characters more approaching those of B. macrophylla, since the leaf-
apex is commonly acuminate and the petioles may be glandular. These branches
flower freely, however, and do not appear to be suckers. The more mature branches
are swollen at the tips with the leaves more densely tufted at the apex. The seed-
lings (Forest Dept. 4352 and 4969) are interesting since the leaves have a long acute
acumen, although they do not appear to be glandular. The cotyledons are about
2-2°5 by 3-4 cm., truncate or very shallowly retuse at the apex and very weakly
cordate to rounded at the base.

The domatia are of unique structure in the genus, although they are of a type
commonly found in Tervminalia. On mature branches they are found usually on
some or most of the leaves, in the axils of the major lateral nerves on the lower surface.
They are often very small, but may be up to about 1:5 mm. across, and are fre-
quently dome-shaped in general outline, with a pore either centrally placed or at the
end of the domatium distal to the nerve axil. The pore is never as broad as the total
diameter of the domatium. In all other known species of Buchenavia domatia, if
present, are simple more or less triangular pocket-like structures formed by the epi-
dermal and cortical layers of the midrib and nerve overarching the nerve axil. In
these cases the pore is thus positioned at and therefore always as wide as the maxi-
mum diameter of the domatium and faces parallel to the leaf surface. As in all
species, the domatia of B. fanshawei are frequently marked by a persistent axillary
tuft of hairs in the young stage. Very often, at an early stage, the domatium is a
simple wart-like object, the pore appearing later by the rupture of the wart. This
is not the case in the other types of domatium (e.g. in B. capitata). In a survey of
domatia in the family Combretaceae, of thirty American species of Terminalia ex-
amined twenty-three were found to lack domatia, three to possess domatia like those
of B. capitata and four to have domatia like those of B. fanshawei. In Buchenavia,
of the twenty-three species available for study (B. discolor being, as stated, unavail-
able) nine (ns. 7-11, 14 and 19-21 of this revision) lack domatia, thirteen (ns. 1-6,
12, 15-17 and 22-24) possess domatia like those of B. capitata and none except
B. fanshawet has the other type.

B. fanshawet is not separable from B. macrophylla on any single character, although
a number of characters are very nearly constant. A combination of characters,
however, will give a good separation in all specimens we have encountered. The
single Brazilian specimen of B. fanshawezi is in fact fairly typical of the species, not
approaching B. macrophylla. It has very conspicuous domatia typical of B. fan-
shawet.

32 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

19. Buchenavia macrophylla Eichl. in Flora 49 : 166 (1866) ; in Mart., Fl. Brasil.
14 (2) : 98 (1867).

Terminalia macrophylla Spruce ex Eichl., loc. cit. (1867), nom. syn.
Buchenavia stellae Cuatrec. in Fieldiana, Bot. 27 (1) : 109 (1950).

Leaf-lamina subcoriaceous to coriaceous at maturity, 3-34 I-5-II cm., (narrowly
elliptic) oblanceolate to narrowly obovate, minutely apiculate to long-acuminate (or
rarely rounded) at the apex with an acumen up to 2 cm. long, acutely to very nar-
rowly cuneate and often markedly narrowed and sometimes decurrent into the
petiole at the base, almost glabrous except pubescent on the midrib and lateral
nerves on both surfaces ; midrib very conspicuous and raised below ; lateral nerves
very conspicuous and raised below, (4) 6-14 pairs ; reticulation rather inconspicu-
ous or fairly conspicuous, not close ; petiole rather slender to stout, very variable in
length, 6-32 mm. long, densely to sparsely pubescent, nearly always conspicuously
biglandular from below the middle to at the top. Peduncle not stout, 6-32 mm.
long, shortly pubescent ; rhachis 20-100 mm. long, densely puberulous ; flowers as in
B. fanshawet. ¥ruit oblong-ellipsoid to ovoid-ellipsoid, terete or very slightly
flattened when fresh, up to 23 X I2 mm., rounded to subacute (apiculate) at the
apex, rounded to very shortly pseudostipitate at the base, very shortly but densely
ferrugineous-tomentellous, indumentum wearing off when very old ; endocarp as in
B. fanshawet.

CoLoMBIA : Caqueta: Florencia, in the cerros La Estrella, 400 m., 30 Mar. 1940,
Cuatrecasas 8863 (US, isotype of B. stellae).

BRAZIL: Amazonas: Near Panuré on Rio Vaupés, Oct. 1852—Jan. 1853, Spruce
2507 in part (BM; BR, lectotype, not seen; K). Cjapo, Rio Vaupés, “ Small
elegant tree 25 ft. Fls. reddish-brown. I was nearly too late for this’, Sept. 1852,
Spruce 2507 in part (K, isolectoparatype). Rio Demeni, tributary of R. Negro,
lake of black water, 12 Oct. 1952, Frdes 28888 (BM). Jaureté, Papury, bank of river,
16 Oct. 1945, Frdes 21156 (K). Mandaos, Pensador, riparian forest of the Igarapé,
to Jan. 1944, Ducke 1542 (K). Mandaos, Pensador, humid spot in non-flooded forest,
20 Aug. 1935 and 3 Feb. 1936, Ducke 35168 (K). Manaos, banks of the Igarapé
Mindu, 8 Sept. 1945, Ducke 1753 (K). Mandaos, along road to Aleixo, 12 Aug.
I Sept. 1936, Krukoff 7921 (BM; K). Humayta, near Tres Casas, basin of Rio
Madeira, on varzea land, 14 Sept.—11 Oct. 1934, Krukoff 6117 (BM; F; K), 6273
(BM; F; K). Parad: Faro, 4 Feb. 1910, Ducke 10649 (BM); same locality,
9 Oct. 1915, Ducke 15798 (BM). Lago de Faro, on banks in rarely flooded places,
22 June 1926, Ducke 13583 (K). Rio Tapajoz, region of the lower waterfalls, 25
June 1918, Ducke 17065 (BM). Rio Sao Manoel, Cachoeira do Calderao, on the
boundary with Mato Grosso State, by Igarapé Preto, 6 Jan. 1952, Pires 3822 (BM).
Rio Pixuna, 40 km. from the mouth of Rio Cupari, between Prainha and Agua Béa,
22 Dec. 1947, Black 47-1964 (BM). Amapd: Rio Araguari, near the Cachoeira do
Paredao, “‘ terra firme’, 25 July 1951, Frées & Black 27695 (BM).

B. macrophylla is apparently the most abundant species of the genus in its area
of distribution. It has not been recorded from Venezuela, although it seems likely
to occur there. It probably also occurs further up the Vaupés in Colombia.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 33

The fruits of B. macrophylla are distinct from those of all other species than
B. fanshawet, the difference between B. macrophylla and that species being discussed
under the latter. In their typical form the leaves are also quite characteristic and
distinctive.

We have little hesitation in including B. stellae, of which we have seen an isotype,
under the present species. It is the only record of B. macrophylla which we know
of from Colombia. All the characters by which B. stellae is supposed to differ
(notably its longer petioles) are found quite frequently in various forms of B. macro-
phylla. Although B. stellae was compared by the author with B. reticulata, it and
B. macrophylla in general are very distinct from that species.

20. Buchenavia longibracteata Fr’es in Bol. Técn. Inst. Agron. Norte 20: 53,
t. 16 (1950).

Leaf-lamina chartaceous at anthesis, 8-23 x 4-8 cm., usually oblanceolate, shortly
acuminate at the apex, narrowly cuneate but not decurrent into the petiole at the
base, almost glabrous above except tomentose on the midrib and densely pubescent
on the lateral nerves, conspicuously ciliate on the margin, sparsely pubescent below
except densely pubescent on the midrib and lateral nerves (at flowering time) ;
midrib conspicuously raised below; lateral nerves conspicuously raised below,
c. I2-16 pairs ; petiole short and stout, c. 6-13 mm. long, shortly pubescent, con-
spicuously biglandular. Rhachis c. 55-110 mm. long, densely pubescent ; flowers
c. 3°5 X 3 mm., with a pubescent ovary but a glabrous upper hypanthium ; bracts
relatively large and conspicuous, foliaceous, c. 7-13 X I-2 mm., with a conspicuous
midrib, pubescent, exceeding the flowers. Fruit so far unknown.

BRAZIL: Amazonas: Cachoeira das Araras, Rio Vaupés (near the Colombian
frontier), ““Arvore, 5 m.”, 1 Nov. 1945, Frdes 21308 (IAN, holotype, not seen; K).

This species is very distinct in its large leafy bracts which, as far as is known, are
unique in the genus. Apart from this character, however, it scarcely seems possible
to distinguish it from B. macrophylla, and indeed it may be an abnormal form of
that species. As the fruits are unknown (the only gathering being in flower) we
prefer to maintain the species for the present. The locality is within the range of
B. macrophylla but we have not seen the slightest trace of large leafy bracts or of a
ciliate leaf-margin in the latter species.

21. Buchenavia pulcherrima Exell & Stace, sp. nov.

Arbor c. 20 m. alta ; ramuli griseo-fusci, fere glabri, apice clavati. Folia spiraliter
ordinata ad ramulorum apices congesta ; lamina subcoriacea, 27:5-29 < 8-10 cm.,
anguste obovata vel oblanceolata, integra, in toto fere glabra, apice abrupte acute
mucronata vel apiculata (apiculo ad 8 mm. longo), basi anguste cuneata, costa media
infra valde prominenti; nervis lateralibus infra prominentibus 9—11-paribus,
venulis conspicuis vix prominentibus; petiolus 30-38 mm. longus, triquetrus,
inconspicuissime glandulosus vel eglandulosus. Inflorescentiae axillares, pendulae,

34. REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

elongato-spicatae ; pedunculus gracilis, c. 30-42 mm. longus, sparse rufo-pubescens ;
rhachis gracilis, c. 70-110 mm. longa, rufo-pubescens. Fructus ellipsoideus, teretius-
culus, immaturus ad 12 x 8 mm., apice basique rotundatus vel latissime obtusus,
pulchre dense rufo-velutinus.

BRITISH GUIANA: Essequibo : 172 km. along Bartica-Potaro road, “60 ft. tree
ro in. diam. basally swollen from Clump Wallaba forest—lvs. tufted to branch ends,
thinly leathery, primary nerves prominent beneath—young fr. on drooping stalks
from below lvs., oval, rusty-velvety pubescent’, 15 Nov. 1943, Field No. F1r485
= Forest Dept. 4221 (K, holotype) ; same locality, “ Seedlings of 422r with coty-
ledons only—perhaps 10 days old, from below parent tree’’, 15 Nov. 1943, Field
No. F1486 = Forest Dept. 4222 (K).

This species is very different from the other two species of the genus known from
British Guiana: B. fanshawei with small leaves and very shortly brownish-pubescent
fruits; and B. megalophylla with large leaves and long-beaked fruits. It is also
distinct from all other members of the genus in its fruits.

The specimen Forest Dept. 4222 in the Kew Herbarium consists of two seedlings
each of which has only a pair of cotyledons on a short hypocotyl. The cotyledons
are very like those of B. fanshawei from the same area: 3:5-3°8 by 4°5-5°5 cm.,
truncate at the apex, obtusely cuneate to almost truncate at the base. We have
not seen cotyledons of any other species of Buchenavia.

The vernacular name for both B. fanshawei and B. pulcherrima is given as
“ Fukadi ”’,

22. Buchenavia huberi Ducke in Bol. Técn. Inst. Agron. Norte 4 : 24 (1945).

Leaf-lamina coriaceous, 2:5-14 Xx I-3-6-7 cm., narrowly obovate or obovate to
elliptic-obovate, rounded or very shallowly retuse or broadly obtuse or very shortly
apiculate (apiculus up to 4 mm. long) at the apex, acutely cuneate at the base and
not or scarcely decurrent into the petiole, almost glabrous except rather sparsely
pubescent on the midrib and lateral nerves on both surfaces ; midrib conspicuous
and raised below ; major lateral nerves conspicuous and raised below, 5—Io pairs ;
reticulation fine but quite conspicuous, close, forming very small inter-reticular
spaces ; petiole distinct and stout, 14-25 mm. long, sparsely pubescent or pubescent,
eglandular or very inconspicuously glandular near the apex. Peduncle slender,
12-24 mm. long, puberulous, indumentum greyish or rufous; rhachis slender, c.
35-95 mm. long, rufous- or cano-puberulous ; flowers with a densely canescent-
sericeous ovary and a glabrous upper hypanthium. Fruit ellipsoid to ellipsoid-
obovoid, more or less terete, 28-31 X 13-14 mm. (c. 20 mm. broad fide Ducke—
fresh material?), abruptly and shortly apiculate (up to 2 mm.) at the apex, shortly
pseudostipitate (2-3 mm.) at the base, cano- or rufous-tomentose at first, indumentum
wearing off but still visible in very old fruits in wrinkles of drupe and at the ends;
endocarp almost terete, slightly longitudinally ridged, c. 25-30 X 12-15 mm.,
subacute at the apex, more or less rounded at the base.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 35

BRAZIL: Amazonas: Manaos, vicinity of the Cachoeira do Mindu, on “ terra

»” ce

firme ”’, “ arvore grande ’”’, 3 Dec. 1943, Ducke 1450 (K ; RB, lectotype, not seen).
Tabatinga, Estrada do Marco, on “ terra firme ’’, ‘‘ arvore muito grande, com sapo-
pemas possantes’’, 26 Nov. 1945, Ducke 1803 (K). Humayta, near Tres Casas,
basin of Rio Madeira, on low terra firma, 14 Sept.—11 Oct. 1934, Krukoff 6472 (BM ;
K). Humayta, near Livramento, on Rio Livramento, basin of Rio Madeira, on

terra firma, 12 Oct._6 Nov. 1934, Krukoff 6794 (BM; F; K).

Ducke also mentions specimens (Ducke 1308 (RB)) collected from the same locality
(same tree?) as the lectotype on 12 Aug. 1943 (subadult flowers) and on 5 Oct. 1943
(young fruits). Apparently the first specimen known was a tree in the gardens of
Belém (Para) Museum (Para State) introduced in 1904 by Dr. J. Huber from the
mid region of the Rio Perus (Amazonas State). There is a specimen from this tree
(Ducke 1281 (RB)) with mature fruit.

This species appears to be very distinct from all others in its characteristic fruits,
perhaps being closest to the glabrous-fruited B. grandis. Very old fruits seem
quite glabrous but on examination remains of the tomentum can be seen in the
wrinkles and at the ends. In the original description Ducke says that the fruits
have a grey tomentum, and this is true of Krukoff’s two gatherings. Ducke’s two
gatherings, however, have a conspicuously rufous tomentum. Such a change
of indumentum from a greyish to a reddish colour during drying is also known in
leaves of Chrysophyllum species. We do not know of it elsewhere in Buchenavia :
B. sericocarpa has retained its greyish-pubescent fruits in the herbarium and B.
pulcherrima is described in a field note as having ‘‘rusty-velvety pubescent ”’
fruits in the fresh state.

23. Buchenavia grandis Ducke in Arch. Jard. Bot. Rio Janeiro 4 : 148 (1925).

Leaf-lamina chartaceous or subcoriaceous, 4°5-I2 X 2-5 cm., obovate to narrowly
obovate, rounded to subacute or rarely apiculate at the apex, narrowly cuneate but
scarcely decurrent into the petiole at the base, almost glabrous (or sometimes very
sparsely pubescent below) except sparsely puberulous to puberulous on the midrib
and lateral nerves on both surfaces ; midrib conspicuous and raised below ; major
lateral nerves fairly conspicuous and somewhat raised below, c. 6-9 pairs ; reticula-
tion rather conspicuous and close, forming fairly small inter-reticular spaces ; petiole
distinct, not stout, (5) 10-20 mm. long, puberulous to sparsely puberulous, eglandular.
Peduncle slender, c. 13-24 mm. long, rufous-puberulous ; rhachis slender, c. 22—100
mm. long, rufous-puberulous; flowers c. 3-4 X 2-3 mm. Ovary glabrous to
densely rufous-puberulous. Distal portion of lower hypanthium glabrous to sparsely
rufous-puberulous ; upper hypanthium glabrous or very sparsely puberulous. Fruit
ellipsoid, slightly 4—5-angled when dry, c. 24-27 X II-I3 mm. (17 mm. wide fide
Ducke), rather gradually acutely apiculate (up to 2 mm.) at the apex, shortly pseudo-
stipitate (c. 2-3 mm.) at the base, glabrous; endocarp (fide Ducke) compressed,
longitudinally ridged, 20-22 * 12-13 mm., 8-10 mm. thick, acute at each end.

36 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

Brazit : Parad: Obidos, 9 Mar. 1909, Ducke 10235 (BM ; MG, lectotype, not seen);
same locality, wood in elevated places, ‘‘ arbor magna fl. viridibus ”’, 15 Sept. 1927,
Ducke 21349 (K). Oriximina, by Rio Trombetas, 5 Feb. 1918, Ducke 16976 (BM,
lectoparatype). In forest in the Quataquara hills near the middle of Rio Tapajoz,
“arbor magna ’”’, 13 Aug. 1923, Ducke 17687 (K, lectoparatype).

Ducke has recorded the species elsewhere in the State of Para: between Mt. and
Lake Curumu (Ducke 17682 (RB)) ; by the Rio Tapajoz waterfalls near Bella Vista
(Ducke 17688 (RB)) ; and in the Serra de Santarém. He has also recorded it from
Maranhao : near Codo on the Rio Itaperurti (Ducke 658 (MG)) ; and near Mirador on
the same river (Lisbéa 2351 (MG)). Besides Ducke’s four gatherings cited above we
have seen no other specimens of this species, although specimens of other species
have been placed here (e.g. of B. pallidovirens and B. hubert).

It is possible that the specimens mentioned represent more than one taxon, al-
though there is no direct evidence for this besides the great variation in pubescence
noted by Ducke. He states that specimens from Tapajoz have glabrous ovaries
and less densely pubescent inflorescences (i.e. rhachides), specimens from Obidos
and Trombetas have moderately pubescent ovaries and inflorescences ; and that
specimens from Maranhao have very densely pubescent ovaries and inflorescences.
We have, however, noted this range of pubescence in B. oxycarpa, where an increased
pubescence is probably correlated with a drier habitat. This is also very likely the
case in B. grandis, as noted by Ducke since he says that trees from the drier regions
(in Maranhao) are smaller (20 m. as opposed to 30-45 m. high). The Maranhao
specimens are also more pubescent.

This is a plant from the lower regions of the Amazon and coastal areas, which may
well account for the fact that we have seen no recent collections of it. It appears
to be most closely related to B. hubert and B. congesta.

Ducke’s specimens 10235 and 16976 (and probably others) were originally deter-
mined (by J. Huber fide Ducke) as Terminalia lucida Hoffmanns. ex Mart., as there
is some resemblance between the leaves of the two species. The fruits, however,
are quite different, those of T. lucida having very broad wings and a dense indumen-
tum. Two gatherings of B. oxycarpa (Krukoff 1241 and 6174) and one of B. fan-
shawei were also originally determined as T. lucida, although the Termuinalia is quite
unrelated and bears only the most superficial resemblance to any species of Buchen-
avia. We do not know of any other Tevminalia species which has been confused
in this way.

24. Buchenavia congesta Ducke in Trop. Woods 90 : 24 (1947).

Leaf-lamina chartaceous or subcoriaceous, 8-15 xX 3-7 cm., obovate or narrowly
obovate, rounded and very sharply and abruptly shortly apiculate (apiculus up to
c. 3 mm.) at the apex, narrowly cuneate and slightly decurrent into the petiole at
the base, very sparsely puberulous to almost glabrous except puberulous on the
midrib and lateral nerves above, sparsely puberulous at flowering time to very
sparsely puberulous or almost glabrous except puberulous on the midrib and lateral

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 37

nerves below at fruiting; midrib conspicuous and raised below; lateral nerves
conspicuous and raised below, slightly raised above, c. 9-13 pairs; reticulation
rather conspicuous, slightly raised below and above, forming small inter-reticular
spaces ; petiole not stout, c. 20-30 mm. long, puberulous or densely puberulous,
eglandular. Peduncle rather slender, 20-35 mm. long, densely rubiginous-puberulous
at flowering time to rather sparsely so in fruit ; rhachis rather slender, c. 35-85 mm.
long, rather densely rubiginous-puberulous ; flowers c. 3-4 xX 2:3 mm. Lower
hypanthium densely rubiginous-puberulous; upper hypanthium rather to very
sparsely rubiginous-puberulous. Fruit oblong-ellipsoid, very slightly flattened to
more or less terete, 22-25 X 9-I2 mm., rounded to broadly obtuse at the apex,
more or less pseudostipitate (pseudostipe under 2 mm. long) at the base, glabrous
from a very young stage.

BRAZIL: Amazonas: Manaos, near Cachoeira do Mindt in a non-flooded wood
in a humid spot, ‘“‘ arbor sat. magna’’ (over 30 m. fide Ducke in descr.), 3 Dec.
1943, Ducke 1465 (F ; RB, lectotype, not seen) ; same locality, 4 Oct. 1946, Ducke
2003 (RB, lectoparatype, not seen) ; same locality, ‘‘ Arvore grande; flér verde
brancacenta da arvore tipica’”’, 25 Sept. 1947, Ducke 2104 (BM).

Apparently only one tree of this species is known and from it all three of the above
gatherings were made. From these we select Ducke 1465 as lectotype since this bears
mature fruits: Ducke 2003 has young fruits and Ducke 2104 (which was not cited
by Ducke) has flowers.

In its densely congested long-petioled leaves and oblong-ellipsoid glabrous fruits
this species appears to be quite distinct from all others, perhaps approaching most
closely B. grandis and to a lesser extent B. macrophylla.

Insufficiently known species
Buchenavia gracilis Glaziou in Mém. Soc. Bot. France 1 (3) : 203 (1908), nom. nud.

Based on Glaziou 5855 from the State of Rio de Janeiro, Brazil. We have exam-
ined the specimen in Herb. Paris and a fragment in the British Museum Her-
barium from the specimen in Herb. Copenhagen. They both consist of young sterile
material alone, which makes their determination quite impossible even to the extent
of specifying Buchenavia or Terminalia as the genus.

Buchenavia macahensis Glaziou in Mém. Soc. Bot. France r (3) : 203 (1908), nom. nud.

Based on Glaziou 18218 from the State of Rio de Janeiro, Brazil. We have ex-
amined the specimen in Herb. Paris and confirmed the genus as Buchenavia. The
specimen consists only of leaves and very young inflorescences and this makes a
comparison with known species extremely difficult. We consider the specimen too
immature to warrant description as a new species or to allow its identification,
although there are a number of similarities with B. viridiflora. A further possibility
is that the inflorescences would have been capitate. A second sheet in Herb. Paris
bears an identical label to the previous specimen, in addition to the correct collector’s
ticket and tag with n. 27125. This specimen is clearly B. tomentosa and was recorded

38 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

as such by Glaziou (loc. cit.). It was doubtless erroneously mounted with a label
of B. macahensis.

Excluded species

Buchenavia fluminensis Glaziou in Mém. Soc. Bot. France 1 (3) : 203 (1908), nom. nud.

Based on Glaziou 6142 from the State of Rio de Janeiro, Brazil. We have ex-
amined the two specimens in Herb. Paris and found them to be an unknown species
of Terminalia.

RAMATUELLA Kunth

RAMATUELLA Kunth, Nov. Gen. & Sp. Pl. 7: 253 (1825) (‘‘ Ramatuela’’).
Type: R. argentea Kunth.

Kunth originally used the spelling Ramatuela, naming the genus after a French
horticulturist whom he called “‘ Ramatuel”’. De Candolle (Prodr. 3 : 16 (1828)
stated that the correct spelling of this Frenchman’s name was Ramatuelle (actually
given as M. de Ramatuelle in a paper published by the latter in Journ. Hist. Nat.
2: 233 (1792)) and thus rightly corrected the spelling of the generic name to
Ramatuella.+

Ramatuella also belongs to the Terminalieae, differing from the other six genera in
the tribe by having 4—5-winged actinomorphic fruits crowded in spherical masses.
It is confined to the headwaters of the Rio Orinoco and Rio Negro and some of their
tributaries in the Amazonas district of Venezuela, the Vaupés district of Colombia
and the Amazonas district of Brazil (see Fig. 1), the whole region consisting of inter-
connecting waterways since the upper Orinoco divides in such a way that some of
its waters, curiously enough, flow into the Amazon via the Rio Negro. The area of
distribution of the genus lies within latitudes 1° S. and 4° N., and occupies only
about 8° of longitude. There appear to be no ecological or geographical differences
between the species.

The genus is closely related to Terminalia and it is highly probable that it was
derived from an original Teyminalia stock. As in Terminalia and Buchenavia
(and indeed all the Tevminalieae) Ramatuella is apetalous. The lower hypanthium
is constricted at the apex but not extended into a long neck as in Buchenavia. It
and the cupuliform upper hypanthium, which bears 4 or 5 triangular calyx-lobes,
are pubescent on the exterior. The 8 or ro stamens are in two whorls, one opposite
to and one alternate with the calyx teeth, and are conspicuously exserted, as is the
style. In bud the stamens are bent double, the anthers being well protected in the
base of the upper hypanthium which possesses a very well-developed hairy disk.
As in Terminalia the anthers are versatile. All species usually have tetramerous
and pentamerous flowers in the same inflorescence. In Ramatuella, as is frequently
the case in Buchenavia, male and bisexual flowers are mixed in the same inflorescence,
in the present genus the former sometimes being the more abundant. The lower

1 Poiret (Dict. Sci. Nat. 44: 427 (1826)) had used the spelling Ramatuella two years previously, but
he gave no reasons for this.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA _ 39

flowers of the inflorescence are more usually male, so that the floriferous rhachis may
be three or more cm. long but give rise to a more or less capitate infructescence.

Ramatuella was surveyed fairly recently by Maguire (in Mem. New York Bot.
Gard. 8 : 130-132 (1953)), who recognized four species, one consisting of two varieties.
The acquisition of further material has now led us to recognize six species and, apart
from the fact that Maguire gave no descriptions, is the only reason for revising the
genus after such a comparatively short interval. These species all consist of small to
medium-sized evergreen trees of riverine forest.

KEY TO THE SPECIES

Lower surface of leaves and fruit densely silvery-sericeous ; wings of fruit short and
broad, very abruptly narrowed at the base into a pseudostipe and at the apex
into a beak 2-4 mm. long ; : 1. R. argentea

Lower surface of leaves for the greater part glabrous or only rather sparsely hairy,
not silvery or sericeous ; fruit various, the wings relatively long ; beak more
or less absent to long :

Fruit under 10 mm. long, deltate-ovate in outline ; wings abruptly narrowed at
the base and apex of the fruit-body ; beak o-5-r mm. long; leaves up to
Poa CMe: : : : 2. R. maguirer

Fruit 10-20 mm. long, iancealave to circular’ in outline, rounded or curved at the
margin ; beak up to 1 mm. long, or 2 or more mm. long and then the wings
gradually narrowing into it:

Leaves glabrous or very sparsely pubescent over the whole of both surfaces ;
fruit acuminate at the apex, beaked, the beak 2-5 mm. long at maturity:

Leaf-lamina 2°5-1I x I-5 cm.; petiole 4-15 mm. long; fruit silvery, ovate
in outline ; beak c. 2-5 mm. long, rather slender ; pseudostipe c. 2 mm.
long. ; ‘ ; : , : . 3. R. virens

Leaf-lamina 8-14 x oy cm. © petiole 20-30 mm. long; fruit fulvous, nar-
rowly ovate to lanceolate ; in outline ; beak c. 2-4 mm. long, rather stout ;
pseudostipe absent ‘ ‘ Fae ce latifolia

Leaves rather densely pubescent at the base of the midrib on the lower surface ;
fruit truncate to obtuse at the apex, beakless or with a beak not exceed-
ing I mm. long:

Leaf-lamina oblanceolate to narrowly oblong ; fruit broadly elliptic to cir-
cular in outline, 12-18 mm. wide, truncate to rounded-retuse at the
apex ; wings markedly crisped, rather thin : . 5.2. crispialata

Leaf-lamina obovate to oblong ; fruit oblong to ovate in outline, 8-15 mm.
wide, obtuse to almost truncate at the apex; wings undulate, rather
thick . : : : : : ; ' . 6. R. obtusa

1. Ramatuella argentea Kunth, Nov. Gen. & Sp. Pl. 7 : 254, t. 656 (1825).—DC.,
Prodr. 3 : 16 (1828).—Eichl. in Mart., Fl. Brasil. 14 (2) : 99, t. 26 fig. 1 (1867).—
Maguire in Mem. New York Bot. Gard. 8 : 131 (1953). (Fig. 4 ¢.)

40 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

Fic. 4. Ramatuella maguirei Exell & Stace (holotype): a, habit (x %); 0, fruit (x2).
R. argentea Kunth (Spruce 3498): c, fruit (x2). R. virens Spruce ex Eichl. (Maguire,

Cowan & Wurdack 30764): d, fruit (x2). R. latifolia Maguire (holotype): e, fruit

(x2). R. obtusa (Maguire) Exell & Stace (holotype): f, fruit (x2). R. crispialata
Ducke (Frdes 28040) : g, fruit (x2).

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 41

Leaf-lamina coriaceous, 2-Io X I-4 cm., oblanceolate to narrowly obovate, retuse
to rounded at the apex, narrowly cuneate and slightly decurrent into the petiole
at the base, almost glabrous above except sparsely or very sparsely puberulous on
the midrib, densely silvery-sericeous below ; midrib conspicuous and raised below ;
major lateral nerves inconspicuous, c. 5—II pairs; reticulation not visible; petiole
usually distinct, 5-16 mm. long, appressed-puberulous. Peduncle (in fruit) rather
slender to stout, 5-35 mm. long, appressed-puberulous ; rhachis up to 6 mm. long.
Fruits crowded into spherical capitula, 8-12 <x 8-13 mm., densely silvery-sericeous ;
pseudostipe stout, abruptly delimited from upper regions, 2-3 mm. long; beak
rather slender, abruptly delimited from lower regions, 2-4 mm. long ; body of fruit
with 4-5 short and broad (1-5 x 3-6 mm.) wings; margin of wings sharply or
bluntly pointed, not undulate.

VENEZUELA or COLOMBIA: Rio Atabapo, Humboldt & Bonpland (P, holotype,
not seen). Rio Guainia, above its confluence with Rio Casiquiare, May 1854,
Spruce 3498 (BM ; K).}

CoLtomMBIA: Vaupés: Rio Atabapo, drowned river margin, frequent on the
Colombian bank between San Fernando and Cacaqual, 130 m., “ bushy tree 7 m. high,
fruit tan’’, 18 Nov. 1953, Maguire, Wurdack & Bunting 36264 (BM). Rio Vaupés
(not seen ; fide Maguire).

BRAZIL: Amazonas: Icana, bank of Rio Cubato, “ Arvore 4 m. com ramagens
densas ”’, 19 Nov. 1945, Frdes 21421 (K).

In its densely silvery-sericeous lower leaf surfaces and characteristic fruits (see
Fig. 4) this species is very distinct from all the other five.

2. Ramatuella maguirei Exell & Stace, sp. nov. (Figs. 4.4, b; 5a, D.)

Arbor parva vel media, c. 20 m. alta ; ramuli griseo-fusci, fere glabri, apice clavati.
Foha spiraliter ordinata ad ramulorum apices congesta, 2-II x I-4 cm.; lamina
subcoriacea, oblanceolata vel elliptico-obovata, apice retusa, basi acuta in petiolum
decurrens, integra, costa infra minute puberula excepta glabra, costa media supra
inconspicua subtus prominenti, nervi lateralibus inconspicuis ; petiolus c. 3-II mm.
longus, sparse puberulus. IJnflorescentiae axillares, spicatae vel capitatae ; pedun-
culus vix validus, 15-50 mm. longus, sparse puberulus ; rhachis ad 33 mm. longa ;
flores masculini et bisexuales in eandem inflorescentiam dispositi. Hypanthium
inferum c. 3-7 X I-2 mm., dense adpresso-pubescens, apice in floribus masculinis
angustatum, in floribus bisexualibus brevissimum et angustissimum; superum
Cc. 3-4 X 3°5-5 mm., adpresso-pubescens vel sparse adpresso-pubescens. Calycis
lobt 4 (vel 52), c. I°5 mm. longi, obtusi. Stamina 8 (vel 10?), exserta ad 6 mm.
Stylus ad c. 5 mm. exsertus, filiformis, glaber. Fvuctus dense congesti, 8-9 x 9-II
mm., (4) 5-alati, rhomboidei vel late ellipsoidei, olivaceo-fusci, minute argenteo-
pubescentes, rostro 0-5-1 mm. longo, alis tenuibus in rostrum stipamque abrupte

1 Both the Rio Atabapo and the Rio Guainia form the Venezuelan-Colombian frontier, and we cannot

be certain to which country these collections belong. No other collection of this species appears to
have been made in Venezuela, although it undoubtedly occurs there.

42 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

contractis amplis lateraliter obtusis vel obtuso-rotundatis margine planis vel leviter
undulatis.

VENZUELA : Amazonas : Alto Rio Orinoco, Caio Yapacana from laguna to mouth,
125 m., “‘ tree 20 m. high, occasional waterside ’’, 17 Mar. 1953, Maguire & Wurdack
34606 (BM, holotype).

BraziL : Amazonas: Rio Curicuriary, tributary of Rio Negro, “ super cataractas,
ad ripas inundatas. Arbor parva fl. albidis”, 23 Nov. 1936, Ducke 34638 (K) ;
same locality, 20 Nov. 1936, Ducke 34639 (K) ; same locality, 22 Feb. 1936, Ducke
34640 (K).

We have pleasure in naming this distinct species, which has been distributed as
R. virens, after one of its collectors, Bassett Maguire, author of a previous paper on
the genus.

R. maguirei is easily distinguished from the other species of the genus by its rather
small almost glabrous leaves and by its fruits, the wings of which are deltate in out-
line and abruptly narrowed above into the very short but distinct beak and below
into a stalk-like base. The fruits are also less pubescent than in any other species
and smaller than in all but R. argentea, to which it is perhaps nost closely related
(except in its lack of sericeous indumentum).

The Venezuelan specimen has mature fruits, while of the three Brazilian specimens
one (Ducke 34640) has mature fruits and all three have open flowers.

There are two distinct types of mature fruit among the material which has hitherto
been identified as R. vivens. These are represented by (a) Maguire, Cowan &
Wurdack 30764, and (b) Ducke 34640 and Maguire & Wurdack 34606. They obviously
belong to two distinct species. We have, however, had great difficulty in deciding
which of them should be linked with Spruce 3758 (the type collection of R. virens)
since the latter has flowers and only very immature fruits. The very young fruits
and ovaries at anthesis give very little clue to their shape at fruiting time. Ducke
34640 has both flowers and fruits and the former differ slightly from the flowers of
R. virens (Spruce 3758), the upper hypanthium being somewhat less pubescent and
the calyx lobes more obtuse (see Fig. 5). Moreover, the very young fruits have
less sinuate wings. We have therefore decided that Maguire, Cowan & Wurdack
30764 belongs to R. virens and that Ducke 34640 and Maguire & Wurdack 34606
represent a new species, described above.

Ducke 34638 and 34639 are interesting in that the rather elongate inflorescences
consist almost entirely of male flowers, the bisexual flowers being confined to one
or two at the extreme apex of the rhachis.

3. Ramatuella virens Spruce ex Eichl. in Mart., Fl. Brasil. 14 (2) : 100, t. 26 fig. 2
(x867).—Maguire in Mem. New York Bot. Gard. 8 : 131 (1953). (Figs. 44; 5¢.)

Leaf-lamina coriaceous, 2°5-II xX I-5 cm., oblanceolate to obovate, retuse to
rounded at the apex, narrowly cuneate and slightly decurrent into the petiole at the
base, almost glabrous on both surfaces except sparsely puberulous on the midrib
below ; midrib conspicuous and raised below ; major lateral nerves inconspicuous,

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 43

Fic. 5. Flowers of Ramatuella. R. maguirei Exell & Stace (Ducke 34638): a, male
flower ; b, bisexual flower. R. vivens Spruce ex Eichl. (type collection) : c, bisexual
flower. R. obtusa (Maguire) Exell & Stace (Frdes & Addison 29040) : d, bisexual flower.
(All x 3.)

c. 8-15 pairs; reticulation not visible; petiole usually fairly distinct, 4-15 mm.
long, sparsely puberulous. Peduncle (in fruit) rather slender, 30~70 mm. long,
appressed-puberulous ; rhachis up to 25 mm. long. Flowers as in B. maguiret
(but see discussion thereunder). Fruits usually crowded into spherical capitula, but
sometimes a few below the head, c. 13-17 X 6-I0 mm., ovate in outline, densely
silvery-sericeous when mature ; pseudostipe abruptly delimited from upper regions,
c. 2 mm. long; beak rather slender, gradually contracted from lower regions, 2—5
mm. long ; body of fruit with 4-5 long and rather narrow (8-12 X 2—4 mm.) wings ;
margin of wings gradually curved, conspicuously but minutely sinuate ; wings rather
thin.

VENEZUELA : Amazonas : Rio Orinoco, Cerro Yapacana, Cano Yapacana, “ river-
ine tree 10-15 m. high, fruit gray-brown, occasional ’’, 6 Jan. 1951, Maguire, Cowan
& Wurdack 30764 (BM; K).

VENEZUELA or COLOMBIA: Rio Guainia, above its confluence with Rio Casi-
quiare, Nov. 1854, Spruce 3758 (type collection; BM; K)!; from type locality,
Schultes & Lopez 9359 (not seen ; fide Maguire).

BraziIL: Amazonas: Rio Negro, Enuixy, Matozinho, 18 May 1947, Froes 22340
(BM).

The problem of the identification of fruiting specimens with Spruce’s type collec-
tion of R. vivens (in flower) is discussed above under R. maguiret.

4. Ramatuella latifolia Maguire in Mem. New York Bot. Gard. 8: 131 (1953).
(Fig. 4 ¢.)
Leaf-lamina coriaceous, 8-14 xX 4-7 cm., oblanceolate to obovate, retuse to
rounded at the apex, cuneate and scarcely decurrent into the petiole at the base,

1 See footnote on p. 41 concerning Spruce 3498 under I. argentea.

44 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

almost glabrous on both surfaces except sparsely puberulous on the midrib below ;
midrib conspicuous and raised below ; major lateral nerves quite conspicuous below,
c. 8-16 pairs ; reticulation visible, irregular ; petiole distinct, (10) 20-30 mm. long,
very sparsely puberulous. Peduncle (in fruit) rather slender, 50~70 mm. long, densely
puberulous ; rhachis up to c.6 mm. long. Fruits crowded into spherical capitula,
14-18 xX 6-IO0 mm., narrowly ovate to lanceolate in outline, densely tawny-puberu-
lous when mature ; pseudostipe absent ; beak rather stout, gradually contracted
from lower regions, 2-4 mm. long; body of fruit with 4-5 long and rather narrow
(10-13 X 2-4 mm.) wings; margin of wings gradually curved, conspicuously but
minutely sinuate ; wings rather thick.

VENEZUELA : Amazonas : 15 km. above San Fernando, Rio Atabapo, Rio Orinoco,
125 m., ‘‘ small tree ca. 10 m. high ’’, “‘ occasional on sandy banks of flood Caatinga ’’,
17 Oct. 1950, Maguire 29258 (NY, holotype).

This species is obviously very closely related to R. virens and may ultimately
even prove to be conspecific. The fruits (see Fig. 4) and leaves, however, show
rather small but distinct differences. The “ cuticular exfoliation ’’ to which Maguire
refers is of very dubious taxonomic value. Whatever is exfoliated (perhaps it is a
surface layer of wax) it is certainly not the leaf-cuticle since in this genus the latter
is extremely thick and extends well down between the epidermal cells.

5. Ramatuella crispialata Ducke in Arch. Inst. Biol. Veg. Rio Janeiro 2 : 65 (1935).

—Maguire in Mem. New York Bot. Gard. 8 : 131 (1953) excl. var. obtusa. (Fig.

4 &.)

Leaf-lamina coriaceous, 6-13 X 2°5~7°5 cm., oblanceolate to narrowly obovate,
retuse at the apex, cuneate but scarcely decurrent into the petiole at the base,
almost glabrous except sparsely puberulous on the midrib above, very sparsely
puberulous to almost glabrous except densely puberulous on the midrib below ;
midrib conspicuous and raised below ; major lateral nerves quite conspicuous below,
c. 8-18 pairs; reticulation visible, irregular; petiole distinct, c. 12-17 mm. long,
puberulous. Peduncle (in fruit) rather slender, 50-95 mm. long, densely puberulous ;
rhachis up toc. 12mm. long. Fruits crowded into spherical capitula, 16-20 « 12-18
mm., broadly elliptic to circular in outline, densely tawny-puberulous when mature ;
pseudostipe absent ; beak absent or up to r mm. long, very abruptly contracted
from the truncate or retuse apex of the fruit ; body of fruit with 4-5 broad and
long (15-20 X 5-7 mm.) wings ; margin of wings rounded, conspicuously crisped ;
wings rather thin.

BraziL: Amazonas: “ Caatinga ad Igarapé Juraxare affl. Rio Vaupés ”’, 2 Nov.
1932, Ducke 25024 (K; RB, holotype, not seen); from type locality, 29 Sept.
1935, Ducke 221 (not seen ; fide Maguire). El Castillo, Rio Negro, Vaupés, Schultes
& Lopez 9298a (not seen ; fide Maguire). Rio Ig¢ana, Urumuturu, mata de caatinga,
“ arvore Io m., recurvando-se sébre 0 rio ’’’, 26 Mar. 1952, Fvdes 28040 (BM).

The large broad-winged fruits of this species render it very easily identifiable.
Even sterile specimens can readily be determined owing to the distinctive shape, size
and pubescence of the leaves.

REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA 45

6. Ramatuella obtusa (Maguire) Exell & Stace, stat. nov. (Figs. 4f; 5d.)
Ramatuella crispialata var. obtusa Maguire in Mem. New York Bot. Gard. 8 : 131 (1953).

Leaf-lamina coriaceous, 3:2-I3 X I-7~-7 cm., narrowly obovate to obovate or
narrowly elliptic to elliptic, retuse to rounded at the apex, cuneate to broadly
cuneate and scarcely decurrent into the petiole at the base, very sparsely puberulous
to glabrous except puberulous to sparsely puberulous on the midrib above, very
sparsely puberulous except densely puberulous on the midrib below ; midrib con-
spicuous and raised below ; major lateral nerves quite conspicuous below, c. 8-18
pairs ; reticulation visible, irregular ; petiole distinct, 7-25 mm. long, densely puberul-
ous. Peduncle (in fruit) rather slender to rather stout, 50-110 mm. long, densely
puberulous ; rhachis up to 15 mm. long ; flowers male and bisexual in the same in-
florescence. Lower hypanthium 4-6 x 2-4 mm., very densely appressed-pubescent,
slightly constricted at the apex; upper hypanthium 3-4°5 x 4-6 mm., densely
appressed-pubescent to more or less tomentose. Calyx lobes 4 (or 5?), c. I°5 mm.
long, obtuse, rather thick. Stamens 8 (or r0?), exserted to 4mm. Style exserted
to 3 mm., stout, densely pubescent for at least the proximal three-quarters. Fruits
usually crowded into spherical capitula but sometimes a few below the head,
10-20 X 8-15 mm., ovate to oblong in outline, densely tawny-puberulous when
mature ; pseudostipe absent ; beak absent or up to I mm. long, abruptly contracted
from the obtuse to almost truncate apex ot the fruit ; body of fruit with 4-5 broad
and long (9-18 x 3-6 mm.) wings; margin of wings rounded, conspicuously un-
dulate ; wings rather thick.

VENEZUELA: Amazonas: Cerro Yapacana, Rio Orinoco, “tree to 7 m. high,
fruit brown, occasional in Caatinga about Yapacana Savanna I.”’, 7 Jan. 1951,
Maguire, Cowan & Wurdack 30796 (NY, holotype).

BRAZIL: Amazonas: Rio Araca, subtributary of Rio Negro, Campinarana,
“ arvore de 2 m., flor esbranquicada, sdlo arenoso ’’, 24 Oct. 1952, Frées & Addison
29040 (BM).

Maguire recognized five taxa (four species, one having two varieties) in the genus
Ramatuella. These fall readily into three easily defined groups :

I—R. argentea, with silvery-sericeous fruits and under surfaces of the leaves, the
fruits with short wings abruptly narrowed at the base and at the apex into a fairly
long beak (Fig. 4c).

2—R. virens and R. latifolia, with nearly glabrous leaves, and fruits with long
wings gradually tapering at the apex into a rather long beak (Fig. 4 d, e).

3—R. crispialata vars. crispialata and obtusa, with leaves nearly glabrous except
for a rather dense brownish indumentum at the base of the midrib on the lower
surface, and fruits with long broad wings abruptly narrowed to a very short beak at
the apex (Fig. 4 f, g).

R. maguirei, our new species, forms a fourth group (see above).

In group 2, Maguire separated R. latifolia, known only from the type specimen,
from R. virens by several small though well-defined characters (see key) which we
regard as sufficient to maintain them as specifically distinct. In group 3, we con-
sider that the differences between R. crispialata var. crispialata and var. obtusa are

BOT; 3; I. 2

46 REVISION OF THE GENERA BUCHENAVIA AND RAMATUELLA

comparable to those between R. latifolia and R. virens and for this reason we have
raised R. crispialata var. obtusa to the rank of species. Maguire himself suggested
that this might ultimately prove to be the best course. We have recently received
a specimen from a second station for R. obtusa, which extends the eastern limit of
Ramatuella to 63° W. This specimen (Frdes & Addison 29040) is of further interest
since it bears flowers and unripe fruits, the type having mature fruits. It has
almost all the characters described by Maguire, except that the leaf base is not
always more obtuse than in R. crispialata, although the leaf is broader in com-
parison with its length. In addition to the characters given by Maguire, the fruit
of R. obtusa is clearly narrower and less truncate at the apex than is that of R. crispi-
alata (see Fig. 4).

Of the six species now recognized, only R. obtusa, R. maguire: and R. virens are
known in flower. The last was figured by Eichler (in Mart., Fl. Brasil. 14 (2) : t. 26
fig. 2 (1867)) but we have re-examined the flowers of this species and compared them
with those of the other two species. The flowers of R. obtusa differ from those of
R. virens and R. maguirei chiefly in their greater amount of indumentum, a feature
also shown by the leaves. This difference is most evident on the outside of the upper
hypanthium and calyx lobes, which are appressed-pubescent in R. virens but densely
appressed-pubescent to more or less tomentose in R. obtusa ; and in the style, which
is entirely glabrous in R. virens but densely pubescent in R. obtusa, at least for the
proximal three-quarters and sometimes along its whole length. Other parts of the
flower are also more pubescent in R. obtusa, but the stamens are completely glabrous.
A further feature is that the style and stamens, which are of the same length, are
somewhat longer in R. vivens, the filaments being over twice as long as the calyx
lobes in the latter species but rarely that length in R. obtusa. This character should
be used, however, with caution, since the filaments and style elongate considerably
at anthesis. The flowers of R. obtusa are also slightly larger, the ovary especially
so, although the latter, of course, soon enlarges after pollination. Furthermore the
ovary is less constricted at the apex in R. obtusa (see Fig. 5).

pe TOGRAMMA
| AND THE IDENTITY OF
__-:- SCHIZOSTAURON

Ro ROSS

pee - BULLETIN OF
i BRITISH MUSEUM (NATURAL HISTORY) —
/ Vol. 3 No.2
“LONDON: 1963 oe

THE DIATOM GENUS CAPARTOGRAMMA
AND THE IDENTITY OF SCHIZOSTAURON

BY

R, ROSS

Php. 47-92 ; 38 Text-figures ; Plates 1-2

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 3 No. 2
LONDON: 1963

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(NATURAL HISTORY), instituted in 1949, is
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THE DIATOM GENUS CAPARTOGRAMMA
AND THE IDENTITY OF SCHIZOSTAURON

By R. ROSS

HusTEDT (in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 757 (1959)) has suggested that
Stauronets crucicula (Grun. ex Cleve) Comber and another species for which he used
the name S. karstenii+ constitute a distinct subgenus of Stauroneis Ehrenb.
characterized by a forking of the central nodule. However, the structure of these
two species is so different from that of the type species of Stauroneis and its near
relatives as to justify treating them, and two hitherto undescribed and closely
related diatoms, as a separate genus. Stauroneis has the central nodule transversely
expanded so that it forms a single band of thickening across the valve, reaching
almost or quite to the margin and interrupting the striae. In the species under
discussion, on the other hand, the valve is convex and the central nodule is connected
to the margin of the valve by two diverging strands of silica on either side. These
are not mere thickenings of the valve surface but are deep and narrow flanges
running between two striae that are scarcely, if at all, more widely separated than
the rest.

Those authors who have treated these species as belonging to a genus distinct
from Stauroneis have referred them to Schizostauron Grun. (Hedwigia 6 : 28 (1867)),
but this name is typified by a totally unrelated species whose correct position is in
Achnanthes Bory. However, the monotypic genus Capartogramma Kuff. (Expl.
Hydrobiol. Lac Tanganyika (1946-1947), Res. Sci. 4 (3) : 27 (1956)) is based on the
species called by Hustedt Stauroneis karstenit. Kufferath’s description shows that
he did not fully understand the structure of the diatom, especially of its valve view,
and he did not realize that it was one already described and figured by O. Miller
(in Engler, Bot. Jahrb. 45 : 88, t. 2 figs. 17, 18 (1910)) and given the name
Schizostauron karstenit by Zanon (Atti R. Accad. Ital., Mem. Sci. Fis. Mat. Nat. 12 :
494 (1941)) ; O. Miiller, although he used that name, did not give it valid publication.
Nevertheless, Kufferath’s description and figures make it impossible to doubt that
his name is based on this species.

When it had been established that Schizostauron was a synonym of Achnanthes,
the correct position and name of all the species that had at any time been referred
to the genus were investigated. In some cases this revealed confusions with species
which have never been placed in Schizostauron. The detailed account of Caparto-
gramma is accordingly followed by a discussion of Schizostauron, including both its
typification and the identity of all the species which have been placed in it, and this
concludes with annotated synonymies of all the species taken into consideration.
Where accurate descriptions are not available in the literature, these have been
given, and in such cases all the specimens I have seen are listed.

1 This is not a validly published name. See under Capartogramma karstenti below (p. 58).

50 CAPARTOGRAMMA AND SCHIZOSTAURON

All the specimens cited in this paper are in the diatom collection of the British
Museum (Natural History), unless there is an indication to the contrary. I am
much indebted to the Director of the Botany Department of the Naturhistorisches
Museum, Vienna, and the Directors of the Botanical and Palaeobotanical Depart-
ments of the Naturhistoriska Riksmuseum, Stockholm, for the loan of specimens
from the collections in their charge, and to Mr. H. S. Job and Mr. Manfred Voigt
for the loan of specimens from their private collections.

CAPARTOGRAMMA Kuf.

CAPARTOGRAMMA Kuff. in Expl. Hydrobiol. Lac Tanganyika (1946-1947), Res. Sci.
4 (3) : 27 (1956).
E Naviculaceis. Valvae symmetricae vel lunatae, convexae, pseudosepto in
utroque polo instructae. Nodulus centralis tigillis binis cum ambobus marginibus
valvae conjunctus ; tigilla quattuor figuram litteram “‘ X ’’ ad instar facientia.

Type : C. jean Kuff. (= C. karstenit (Zanon) Ross).

The characteristic feature of this genus is the form of its central nodule and
associated structures. The nodule is rectangular, its longer axis being transapical
and with a length of one-fifth to one-third of the width of the valve. From each
corner a strut of silica runs to the margin of the valve, the four struts forming a
figure in the shape of the letter ““X”’. These struts are flattened, ribbon-like
strands of silica and one edge is attached to the inner surface of the valve. The
cross-section of these struts is curved, at least near the margin of the valve, and the
inner, or free, edge is turned towards the apex of the valve ; this edge is continuous
with an inwardly projecting flange that runs along the valve margin and connects
with the pseudoseptum. The term used in the latin diagnoses for these struts is
tigillum (= a little beam). One or more of the struts is at times duplicated or
forked, a single row of puncta, rarely more, lying between the two forks. This
frequently occurs in the smallest specimens of C. crucicula (Grun. ex Cleve) Ross
(Plate 1 B) and on the ventral side in C. amphoroides Ross (Fig. 14), and also, but
much more rarely, in C. karstenit and on the dorsal side in C. amphorotdes.

The struts and pseudosepta of Capartogramma are laid down subsequently to
the formation of the rest of the valve. In dividing frustules containing young
valves at the appropriate stage, these can be seen to have the raphe and striae
completely formed but no trace of the tigilla or pseudosepta. It is possible in such
specimens to see that there is no appreciably larger gap between the striae where
the tigilla will form.

Using the light microscope it was not possible to be sure of the detailed form of
the struts, or whether they were attached to the valve surface throughout their
length. The description given above is based on stereoscopic electron micrographs,
for which I am much indebted to Dr. K. Little of the Nuffield Orthopaedic Centre,
Oxford. C. karstenii, C. crucicula and C. amphoroides were investigated by this
technique (Plate 1) ; C. ryhombicum Ross, which is much rarer in the material available

CAPARTOGRAMMA AND SCHIZOSTAURON 51

to me, shows such great similarity to C. karstenit and C. crucicula under the light
microscope that there can be little doubt that its structure is identical. C.
amphorotdes differs somewhat from the other three species, all of which are sym-
metrical about the apical axis.. It, on the other hand, has the form characteristic
of the genus Amphora Ehrenb. ex Kiitz.; the valves are lunate, with a convex
dorsal margin and an almost straight ventral one, and the girdle is wider on the
dorsal side. The struts on the dorsal side are not very divergent and the electron
micrographs show that they are very similar to those of the other species ; those on
the ventral side are more divergent and are much more expanded parallel to the
valve surface, leaving only a small gap between the two at the centre. It is clear,
however, that the differences between this species and the other three are no more
than minor modifications of the same plan. The fact that these four species all
clearly belong to the same genus raises the question of the taxonomic status of
Amphora and supports the view, tentatively put forward by Cleve in his Synopsis
of the Naviculoid Diatoms (K. Svensk. Vetensk.-Akad. Handl., Ny Féljd, 27 (3) : 99
(1896')), that the various subdivisions of that genus are more closely related to
symmetrical forms of similar structure than they are to one another.

The electron micrographs also provide information about the structure of the
striate part of the valve. Between each pair of striae there is a low rounded ridge
of thickening on the inner surface of the valve. Each stria consists of a single row
of holes (alveoli) through the main substance of the valve ; these are broadly
elliptic to circular, the long axis of the elliptic ones being across the striae, i.e.
parallel to the raphe (Plate 2 ; Fig.1 a). On the outer side of the valve each alveolus
opens through a bar-shaped slit, which is normally parallel to the raphe and is
about one and a half times as long as the alveolus and one-third as broad as it.
These slits are not formed in a separate membrane but in the outer layer of the main
membrane of the valve (Plate 2c). On the inner side of the valve a fine membrane
with pores in triangular tesselation stretches across each alveolus. As the walls
slope inwards for a short distance from this surface, the area covered by the membrane
is somewhat larger than the minimum section of the alveoli. In all three species
examined the centre-to-centre spacing of the pores in this fine membrane is c. 150 A.
Alveoli of similar structure occur in some species of Navicula sect. Lineolatae, e.g.
N. cryptocephala Kitz. (Helmcke & Krieger, Diatomeenschalen in Elektronen-
mikroskopischen Bild 1 : t. 69 (1953)), and in Scoliopleura tumida (Bréb. ex Kiitz.)
Rabenh. (Helmcke & Krieger, op. cit. 2 : t. 177 (1954), whilst those of Pleurosigma
angulatum (Quekett) W. Smith (Helmcke & Krieger, op. cit. 1 : tt. 61-64 (1953))
and other species of that genus are of the same general plan ; the spacing of the
fine pores in the inner membranes of all these species is also of the same order,
from about 135 A. to about 180 A. In the specimens of Capartogramma examined

1 Although the title-page of this work is dated 1895, it does not appear to have been published before
1896. A copy was received by the Royal Microscopical Society between 15 April and 20 May 1896
(Journ. R. Microsc. Soc. 1896 : 368 (30 June 1896)), and it was not noticed elsewhere until later, viz.:
Nat. Nov. 18 : 326 (second half of June 1896) ; Nuova Notarisia 7 : 94 (July 1896) ; Hedwigia 35 :
(104) (30 October 1896). It is also clear that Schiitt had not seen a copy when preparing his account
of the Bacillariaceae for Engler and Prantl’s Natiirlichen Pflanzenfamilien (1 (1, b) “ Gedruckt im Juli
1896’; cf. p. 33), and Van Heurck had only seen a manuscript when writing his Treatise on the
Diatomaceae (cf. pp.105, 125), the preface of which is dated 19th September 1896.

52 CAPARTOGRAMMA AND SCHIZOSTAURON

there seems, however, to be another thin amorphous membrane within that in
which the alveoli lie ; this seems the only explanation for the fact that a small
number of alveoli in which both the bar-shaped slit and the pattern of fine pores
can be seen are appreciably more electron-transparent than the remainder (Plate 2 A).
No such membrane has been reported previously in any diatom and it may be that
it is not part of the silica valve but the consequence of imperfect cleaning or washing.
This seems unlikely, however, for the specimens were cleaned by boiling in a mixture
of concentrated hydrochloric and nitric acids, followed by boiling in concentrated
sulphuric acid into which crystals of potassium chlorate were sprinkled, and were
washed in dilute ammonia and at least seven changes of distilled water.

(i) (ii)

Fic. 1. Diagrammatic cross-sections through alveoli at right-angles to (i) and along the
line of (ii) the striae : a, Capartogramma crucicula (Grun. ex Cleve) Ross ; b, Stauroneis
smithit Grun.; c, S. anceps Ehrenb. (All x 50,000; the horizontal scale exact but
the vertical scale only approximate.)

Cleve (K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 26 (2) : 141-143 (1894)), in
his discussion of the classification and affinities of Navicula sect. Microstigmaticae,
in which he placed Capartogramma crucicula and all the species of Stauroneis then
known, attached considerable importance to the presence or absence of intercalary
bands in the girdle and used their presence as the character distinguishing one
of the groups into which he suggested the section should be divided. It is of interest,
therefore, to note that intercalary bands are present in C. karsteni, C. rhombicum
and C. amphoroides but not in C. crucicula. The close resemblance between these
species in all other respects makes it clear that here at least this character is not
of importance above the specific level.

Since the two species of Capartogramma already known have been treated in

CAPARTOGRAMMA AND SCHIZOSTAURON 53

the past as belonging to or being closely related to Stauroneis, stereoscopic electron
micrographs were taken of four species of that genus : S. phoenicenteron (Nitzsch)
Ehrenb., which is the type species, S. anceps Ehrenb., S. smithii Grun., and S. acuta
W.Smith. The specimens of the first three all came from a gathering from Western
Wood, Cheldon, Devon, collected and cleaned by the late R. F. Bastow, and those
of the last from Petersdorf, near Gleiwitz, Silesia (now Gliwice, Poland), the gathering
distributed as Rabenh., Alg. Mitteleur. n. 847. The last two of these species
resemble Capartogramma in having pseudosepta at the apices, whilst the other
two do not possess these.

The stauros of Stauroneis phoenicenteron, S. anceps and S. acuta is as broad as
the central nodule is long and widens outwards. It is an area of moderate thickening,
not projecting further into the interior of the frustule than the central nodule itself,
and in S. anceps it becomes thinner as it approaches the margin, often so much so
that there are short striae opposite it at the margin. In S. smithii, on the other
hand, the stauros is much narrower than the length of the central nodule ; the
width of the gap between the striae on either side of it is only twice that between
the other striae. It is also very deep, projecting much further into the interior
of the frustule than the central nodule does, and near the margin its inner edge
expands and continues as a flange along the valve margin. At the poles this flange
widens to form the pseudoseptum. S. smithii thus resembles a species of Caparto-
gramma with the two tigilla on either side fused into one.

Stauroneis smith resembles Capartogramma more closely than it does S.
phoenicenteron, S. anceps and S. acuta in the fine structure of the valve also. Its
striae, like those of Capartogramma, are separated by a rounded ridge and the
individual alveoli are more or less circular (Fig. 1 6). They open on the outer
side through broadly elliptic pores, whose long axis is across the striae, i.e. parallel
to the raphe, and is somewhat longer than the diameter of the alveoli, whilst their
short axis is about three-quarters of that diameter. On the inner side of the alveoli
there is a fine membrane with pores in triangular tesselation, but no indication of
an amorphous third membrane was seen in this species.

In Stauronets phoenicenteron, S. anceps and S. acuta the striae are sunk into the
inner surface of the valve in steep-sided troughs and consist of elliptic alveoli
whose long axis is in the direction of the striae (Fig. I c). The distance between
the alveoli is only about half the length of the alveoli themselves, and about half-way
between each pair a narrow flange runs across the trough in which the striae lie.
The alveoli open to the outside through broad slits parallel to the direction of the
striae ; these slits have rounded ends and parallel sides, and are almost as long as
the alveoli and one-half to one-third as wide as them. There is a fine membrane
with pores in triangular tesselation on the inner side of the alveoli stretching from
the tops of the flanges across the trough and thus covering all of this. No trace of
an amorphous membrane was seen.

Stauroneis smithii, as will be appreciated, resembles Capartogramma much more
closely than it does the other species of Stauroneis examined. In S. smuthit var.
sagitta (Cleve) Hust., of which no unmounted specimens for examination with the

54 CAPARTOGRAMMA AND SCHIZOSTAURON

electron microscope were available, the stauros on one or both sides is often divided
so that it forms two parallel bars of thickening with a single stria between them.
As far as can be seen with the light microscope the stauros is deeper than wide,
as in the specimens of S. smithit var. smithit examined under the electron microscope,
and these divided arms of the stauros thus resemble closely the divided tigilla
sometimes found in Capartogramma and especially frequent in specimens of C.
crucicula from Lake Tanganyika.

There are various other species of Stawroneis whose appearance under the light
microscope suggests that in them the stauros is a deep and narrow bar of thickening
rather than a broad area of not very deep thickening such as is found in S. phoenicen-
teron,S. anceps and S. acuta. It is probable, therefore, that when it has been possible
to study the majority of the many species of the genus with the electron microscope
it will be found necessary to divide it. This division, however, will not be that
originally suggested by Cleve (K. Svensk. Vetensk.-Akad. Handl., Ny Féljd, 26 (2) :
141 (1894)) into those with and those without pseudosepta, for S. acuta has pseudo-
septa whilst S. phoenicenteron and S. anceps do not, but both the stauros and the fine
structure of all three are the same. If and when sufficient information to justify a
division of the genus Stauroneis has been accumulated, it will be possible to decide
whether S. smithit and its allies should be treated as a separate genus closely related
to Capartogramma or included within that genus. Meanwhile it seems best to
restrict Capartogramma to those species with two tigilla on either side of the central
nodule and not to transfer Stauroneis smithii from its present position.

KEY TO THE SPECIES

Frustules symmetric about the apical axis ; valves rhombic lanceolate or elliptic :
Valves strongly convex, about 4 deep, apices narrow, long-rostrate ; striae not
more than 17 in I0y, parallel or convergent at the apices ; pseudosepta

markedly inclined to the apical plane, as long as the rostrate apices
1. C. karstentw
Valves not strongly convex, about 2yu deep, with apices not produced, subrostrate,
or rostrate and broad ; striae radiate throughout or 20 or more in I0u ;
pseudosepta almost parallel to the valvar plane, much shorter than the apices

when these are rostrate :

Valves rhombic or lanceolate, apices acute, sometimes subrostrate ; striae
radiate throughout ; pseudosepta 2:°5u long or more . 2. C. rhombicum
Valves elliptic or oblong, apices broad, rostrate, subrostrate or not produced ;
striae parallel at the apices ; pseudosepta 2u long or less . 3. C. crucicula
Frustules asymmetric about the apical axis ; valveslunate . 4. C. amphoroides

1. Capartogramma karstenii (Zanon) Ross, comb. nov. (Plates I A, 2 B, C;
Figs. 2, 3.)

Schizostauron crucicula sensu G. S. West in Journ. Linn. Soc. Lond., Bot. 38 : 156, t. 8
fig. 12 (1907) pro parte ; non Cleve.

CAPARTOGRAMMA AND SCHIZOSTAURON 55

[Navicula sect.] Schizostauron karstenit O. Muell. in Engler, Bot. Jahrb. 45 : 88, t. 2 figs.
17, 18 (1910)!; non Navicula karstenii Pant. (1902).

Schizostauron karstenit Zanon in Atti R. Accad. Ital., Mem. Sci. Fis. Mat. Nat. 12 : 494,
t. 2 fig. 4 (1941).—Monteiro in Estud. Ens. Doc. Junta Invest. Ultramar 72 : 34, t. 1
fig. 7 (1960).

Capartogramma jeanii Kuff. in Expl. Hydrobiol. Lac Tanganyika (1946-1947), Res. Sci.
4 (3) : 27, t. 1 fig. 7 (1956).

Stauroneis karstenit Hust. in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 757, fig. 1117 (1959),
nom, nud.

Frustulum rectangulare, angulis rotundatis, cingulo e copulis composito. Valva
35-60 longa, 9-13 lata, valide convexa, c. 4u alta, anguste elliptica, marginibus
plerumque leniter triundulatis, rostrata, apicibus angustis longis ad finem truncatis.
Pseudosepta c. 6y longa, in planitiem valvarem inclinata, marginibus interioribus
crassis. Raphe filiformis, leniter undulata, poris centralibus distantioribus, fissis
terminalibus in idem latus inclinatis. Area axialis angusta ; area centralis circularis.
Noduli terminales parvi ; nodulus centralis rectangularis. Tigilla duo in altero
latere inter se angulum 30°-60° facientia. Striae leniter radiatae, in apicibus
parallelae convergentesve, tenues, II-17 in Iow ; punctac. 25 in I0uw.

In tropical African fresh waters ; recent from Lakes Victoria, Tanganyika, Zwai
(fide Zanon), Nyassa (fide O. Miller and Monteiro) and Malomba (fide O. Miiller),
and in a stream near Utengule, Tanganyika (fide O. Miiller) ; fossil in a pleistocene
deposit near Bularaba, Nigeria.

UcanDA : Buganda : Lake Victoria, Buvuma Channel, S. end (0° 17’ N., 33° 10’
E.), bottom mud, 30 m., 10 Oct. 1952, Ross 1156 (B.M. 71996). Lake Victoria,
Buvuma Channel, N. end (0° 23’ N., 33° 18’ E.), bottom mud, 22 m., 10 Oct. 1952,
Ross 1158 (B.M. 71999).

TANGANYIKA : Western Province: Lake Tanganyika, near Kirando, plankton,
1 Dec. 1904, Cunnington 175 (B.M. 34180). Lake Tanganyika, Kala Langabo Bay,
near Kigoma, on stone in water c. 50 cm. deep, 30 Jan. 1953, Ross 1457 (B.M. 72100) ;
same locality, on stems and adventitious roots of Phragmites mauritianus, 30 Jan.
1953, Ross 1458 (B.M. 72102). Lake Tanganyika, Kigoma, on submerged leaves of
Paspalidium geminatum, 31 Jan. 1953, Ross 1468 (B.M. 72108) ; same locality, on
Potamogeton pectinatus, 31 Jan. 1953, Ross 1470 (B.M. 72110, 72111, 72116, 72117) ;
same locality, on Lagarosiphon sp. floating in harbour, 31 Jan. 1953, Ross 1473
(B.M. 72129) ; same locality, on roots of Phragmites mauritianus c. I m. deep,
31 Jan. 1953, Ross 1474 (B.M. 72131) ; same locality, on Ceratophyllum demersum
in harbour, 31 Jan. 1953, Ross 1477 (B.M. 72137) ; same locality, scrapings from
jetty wall, 31 Jan. 1953, Ross 1478 (B.M. 72139) ; same locality, bottom mud in
harbour, 2 Feb. 1953, Ross 1512 (B.M. 72141, 72142). Lake Tanganyika, c. 1 km.
off mouth of Luichi River, on shells brought up in trawl, 3 Feb. 1953, Ross 1531
(B.M. 72143) ; same locality, on roots brought up in trawl, 3 Feb. 1953, Ross 1532
(B.M. 72145) ; same locality, mud from large mussel shell brought up in trawl,
3 Feb. 1953, Ross 1533 (B.M. 72147). Lake Tanganyika, c. 1-5 km. S.W. of Ujiji,
bottom mud, 80 m., 3 Feb. 1953, Ross 1535 (B.M. 72149, 72150). Lake Tanganyika,
c. i km. off mouth of Luichi River, bottom mud, 15 m., 3 Feb. 1953, Ross 1536

1This name is not validly published. (see p. 57).

56 CAPARTOGRAMMA AND SCHIZOSTAURON

(B.M. 72151). Lake Tanganyika, Kigoma, scrapings from hull of M.V. “ Sangala ”’,
31 Jan. 1953, Ross 1537 (B.M. 72153). Lake Tanganyika, c. 500 m. offshore between
Ujiji and Bangwe Point, on various weeds tangled in trawl from c. 8 m. deep,
4 Feb. 1953, Ross 1539 (B.M. 72157, 72158) ; same locality, tangled in meshes of
trawl from c. 8 m. deep, 4 Feb. 1953, Ross 1540 (B.M. 72150). Lake Tanganyika,
Bangwe Point, on rocks just below water level, but exposed during moderate swell,
4 Feb. 1953, Ross 1541 (B.M. 72161) ; same locality, pool c. 30 cm. above water
level, often entered by swell, 4 Feb. 1953, Ross 1543 (B.M. 72166). Lake Tanganyika,
Kigoma, bottom sand, c. I°5 m., 4 Feb. 1953, Ross 1549 (B.M. 72171, 72172, 72173,

72174).

Fics. 2-3. Capartogramma karstenti (Zanon) Ross. Fig. 2, valve view (B.M. 72116,
specim. n. 1); Fig. 3, girdle view (B.M. 72116, specim. n. 5). (Both x 1,000.)

FOSSIL: Pleistocene :

NIGERIA : Bornu Province: c. 7 km. N. of Bularaba (11° 04’ N., 11° 48’ E.),
diatomite, Nigerian Geol. Survey P.433 (B.M. 36431, 36432).

O. Miiller’s and Monteiro’s records of this species from Lake Nyassa, O. Miiller’s
from the shallow Lake Malomba, and G. S. West’s from Lake Tanganyika are based
on specimens from plankton hauls made close inshore. Zanon reported the species
as being frequent in gatherings of bottom mud from Lake Zway, and Kufferath
found it in bottom mud from Lake Tanganyika. It was rare in the two of my
gatherings from Lake Victoria in which it occurred ; both these were bottom muds.
In my gatherings from Lake Tanganyika it was commoner in bottom samples than
in epiphytic ones. It thus seems probable that it is primarily a free-living benthic
form, occurring not only on the bottom but also in the films of diatoms and other
algae growing as epiphytes on higher plants, but that it is, in addition, a facultative
plankter, surviving and multiplying in the plankton when lifted into the water mass
by wave action or similar causes. The species composition of the Nigerian diatomite
in which it occurs indicates deposition close inshore in a freshwater lake.

CAPARTOGRAMMA AND SCHIZOSTAURON 57

The specimens I have seen and the published accounts of the species indicate
that it varies very little, but occasionally one of the tigilla is forked. O. Miiller,
at the conclusion of his account of it, distinguished a variety in which the longitudinal
margins of the frustule when seen in girdle view are convex, not straight. The size
range of the specimens he referred to this variety is near the lower limit for the
species, and this convexity is possibly a characteristic of small cells and, as such,
not a justifiable basis for a taxonomic distinction. There is also the possibility
that this variety was based on frustules of C. amphoroides. Careful examination is
often necessary to distinguish large frustules of that species in which the longitudinal
margins are scarcely convex (see Fig. 13) from those of C. karstenii, and if it were
rare in the samples O. Miiller examined and he only saw whole frustules in girdle
view, he might have failed to realize that he was dealing with a distinct species.

Neither in G. S. West’s collection nor among the slides he sent to the British
Museum (Natural History) on completion of his report on the algae collected by
Cunnington on the Third Tanganyika Expedition is there any material of Cunnington
81 or Cunnington 135. There is, however, a preparation of cleaned diatoms from
Cunnington 175, the third of the gatherings from which West reported Schizostauron
crucicula. This contains Capartogramma karstenit and not C. crucicula. West’s
figure, also, is clearly of C. karstenii ; the dimensions he gives and his description
of the outline both fit C. karsteniz, but his value for the closeness of the striae is that
characteristic of C. crucicula. He shows no striae in his figure and one is left to
wonder whether C. crucicula was present in one of the gatherings cited or whether
his value for striation density is an error. (Was it copied from Cleve’s account of
C. crucicula in the Synopsis of the Naviculoid Diatoms ?)

O. Miiller’s description of the species is a full and accurate account of it and
appears under the name Schizostauron karstenit. Above it is the heading “‘ Sectio
Schizostauron Cl. N. D. I. p. 141’, and the typography of this indicates that it is a
subsidiary heading under the main heading “ Naviculae Microstigmaticae Cl.”
O. Miller, in his treatment of Naviculaceae, followed that in Cleve’s Synopsis of the
Naviculoid Diatoms (for which see pp. 63-64 below). It is clear that he cannot be con-
sidered as having validly published the specific name Schizostauron karstenii, for
he did not regard Schizostauron as the name of a genus, and “ the name of a species
is a binary combination consisting of the name of the genus followed by a single
specific epithet ” (Art. 23 of the International Code of Botanical Nomenclature, 1961
edition). Neither can he be held to have published the name Navicula (sect.
Schizostauron) karstenii for he did not definitely indicate that the specific epithet
was to be used in that particular combination (cf. Art. 33 of the International
Code) ; there is no reason for supposing that he was not aware of the previous
publication of Navicula karstenii Pant. for a different species.

Mills (Index Gen. & Sp. Diat.: 1438 (1934)) lists the name “ Schizostauron
Karsteni”’, but with a query and in the type that indicates that the name is a
synonym. Therefore he cannot be regarded as accepting the species and accordingly
he did not give the name valid publication, in spite of his recognition of Schizostauron
as a genus and his citation of O. Miiller’s description. Zanon, in 1941, accepted

58 CAPARTOGRAMMA AND SCHIZOSTAURON

Schizostauron as a genus and S. karstenii as a distinct species and gave the latter
name valid publication by reference to O. Miiller’s description. Hustedt in 1959
used the name Stauroneis karstenit in discussion and gave a figure of the species
but no description nor any reference to O. Miiller’s description or Zanon’s valid
publication of the name; he therefore did not give the name Stauronets karstenii valid
publication either as a new combination or as an independently published new name.

Kufferath’s account of his Capartogramma jeanii is very inadequate. He
apparently worked with fluid mounts and only saw whole frustules. In consequence
he had great difficulty in getting a valve view. His drawing of this aspect of the
diatom shows apices that are much too broad for C. karstenii and his representation
of the central nodule and struts is no more than a caricature. Nevertheless the
slope of the pseudosepta and the depth of the valve shown in his figure of the girdle
view of a frustule show clearly that his name was applied to this species.

2. Capartogramma rhombicum Koss, sp. nov. (Figs. 4-7.)

Frustulum rectangulare, angulis rotundatis, cingulo e copulis composito. Valva
20-55y. longa, 4:5-I2°5u lata, plana (parte prope marginem excepta), c. 2y alta,
lanceolata, anguste lanceolata vel rhombica, aliquando subrostrata, apicibus
angustis rotundatis. Pseudosepta 2:5-5u longa, planitie valvari fere parallela,
marginibus interioribus paullo incrassatis. Raphe filiformis, fere recta, poris
centralibus approximatis, fissis terminalibus in idem latus inclinatis. Area axialis
angustissima ; area centralis rectangularis, parva. Noduli terminales parvi ;
nodulus centralis rectangularis, parvus. Tigilla duo in altero latere inter se angulum
30°-60° facientia. Striae leniter radiatae, 15-22 in 10yu, dense punctatae. |

Fics. 4-7. Capartogramma rhombicum Ross. Fig. 4, valve view (holotype) ; Fig. 5,
valve view (B.M. 72115, specim. n. 4) ; Fig. 6, girdle view (B.M. 72115, specim. n. 5) ;
Fig. 7, valve view (in B.M. 72143). (All x 1,000.)

Holotype : specimen no. 2! in praep. n. B.M. 72114 in Coll. Diat. Herb. Mus. Brit.

From the shores of Lake Tanganyika near Kigoma.

1 The second specimen from the left-hand end of the row as seen under the microscope, the top of
the labels on the slide being to the left.

CAPARTOGRAMMA AND SCHIZOSTAURON 59

TANGANYIKA : Western Province: Lake Tanganyika, Kigoma, on Potamogeton
pectinatus, 31 Jan. 1953, Ross 1470 (B.M. 72110, 72111, 72114, 72115) ; same
locality, on Lagarosiphon sp. floating in harbour, 31 Jan. 1953, Ross 1473 (B.M.
72129) ; same locality, on Ceratophyllum demersum in harbour, 31 Jan. 1953,
Ross 1477 (B.M. 72137) ; same locality, bottom mud in harbour, 2 Feb. 1953,
Ross 1512 (B.M. 72175). Lake Tanganyika, c. 1 km. off mouth of Luichi River,
on shells brought up in trawl, 3 Feb. 1953, Ross 153r (B.M. 72143).

This species has so far been found only in five of my gatherings from Lake
Tanganyika. Three of these were of epiphytes from flowering plants in the harbour
at Kigoma, the fourth of bottom mud at the same place, and the fifth of a growth
of algae and diatoms on empty shells of a freshwater snail brought up in the trawl
from a haul made about 1 km. off the mouth of the Luichi River. This suggests
that the characteristic habitat of the species is the algal felt that grows on solid
substrata in shallow water. It is probably free-living in this habitat and not
attached.

The specimens from Kigoma harbour (Figs. 4-6) and those from off the mouth
of the Luichi River (Fig. 7) clearly belong to two different populations. The
latter are smaller (20-32u long as against 27-55u) and more slender (4:5—6yu
broad as against 9-12), especially when compared with the shorter specimens
from Kigoma harbour ; they have more radiate striae and more divergent tigilla.
More material is necessary, however, before one can decide whether these differences
are indicative of infraspecific taxonomic separation.

Within the population from Kigoma harbour there are differences correlated with
total size. The largest specimens (Fig. 4) have a rhombic outline with acute
apices that are not produced, and the pseudosepta are 4—5u long, whilst the
smaller specimens (Fig. 5) have a lanceolate outline, subrostrate apices, and shorter
pseudosepta only about 2-5y long ; also the tigilla of the larger forms are much
broader parallel to the valvar plane. The variation in these characters is, however,
clearly continuous. That in outline is of interest in that it is unusual for produced
apices to be characteristic of the smaller diatoms in a population when the apices
of the larger ones are not produced.

3. Capartogramma crucicula (Grun. ex Cleve) Ross, comb. nov. (Plates I B,
2A; Figs. I a, 8-11.)

Schizostauron crucicula Grun. ex Dickie in Journ. Linn. Soc. Lond., Bot. 18 : 131 (1880),
nom. nud.

Schizostauron crucicula Grun. ex Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Foljd,
18 (5) : 16, t. 3 fig. 44 (1881).—De Toni, Syll. Alg. 2 : 225 (1891).—Zanon in Comment.
Pont. Acad. Sci. 5 : 55, t. 2 fig. 13 (1941).—Freng. in Rev. Mus. La Plata, Nueva Ser.,
Bot. 3 : 245, t. 1 fig. 19: (1941); op. cit. 8: 83; °t. 1 fig. 31 (1953).

Stauroneis crucicula (Grun. ex Cleve) Cleve, op. cit. 26 (2) : 151 (1894).?

Stauroneis crucicula (Grun. ex Cleve) Comber in Cat. Afr. Pl. Welw. 2 : 385 (1901) excl.
specim.; non S. crucicula W. Smith (1853).—Boyer, Diat. Philad. : 89, t. 27 fig. 10 (1916):
in Proc. Acad. Nat. Sci. Philad. 79, Suppl.: 426 (1928).—Hust. in Hedwigia 63 : 157

1 This name, although used here by Cleve, was not validly published by him (see pp. 63-64).

60 CAPARTOGRAMMA AND SCHIZOSTAURON

(1921) ; in Gessner & Vareschi, Ergebn. Deutsch. Limnol. Venezuela-Exped. 1952 I : 114
(1956).—Krasske in Arch. Hydrobiol. 35 : 555, t. 19 fig. 10 (1939).—Choln. in Osterr.
Bot. Zeitschr. 103 : 87, figs. 128-130 (1956) ; op. cit. 104 : 83, fig. 271 (1957) ; in
Portug. Acta Biol. 6 : 82, t. 4 figs. 123-126 (1958) ; tom. cit.: 138, t. 7 fig. 162 (1958) ;
in Nova Hedwigia 2: 115, t. 9 figs. 334, 335 (1960).—Woodhead & Tweed in Hydro-
biologia 11 : 380 (1958) ; op. cit. 15 : 279 (1960).—Round in Journ. R. Microsc. Soc.
80 : 64, fig. 33 (1961).—Mbodlder in Bull. Comm. Géol. Finl. 198 : 37, t. 1 figs. 7, 13 (1962).

[Navicula sect.] Schizostauron crucicula O. Muell. in Engler, Bot. Jahrb. 45 : 88, t. 1 figs.
14-16 (1910).?

Schizostauron brasiliense Zimmerm. in Broteria, Bot. 16 : 86, t. 2 fig. 3 (1918).

Stauroneis merrimacensis Woodhead & Tweed in Rev. Algol., Nouv. Ser., 5 : 143, t. 15
fig. 27 (1960).—Reimer in Proc. Acad. Nat. Sci. Philad. 113 : 199, t. 2 fig. 6 (1961).—
Choln. in Revista Biol. 3 : 69 (1962).

Stauroneis grunowii sensu Choln. in Hydrobiologia 19 : 104 (1962) ; non Rabenh.

Frustulum rectangulare, angulis rotundatis, cingulo sine copulis. Valva 10-35
longa, 4°5-gy. lata, plana (parte prope marginem excepta), c. 2y alta, elliptica vel
oblonga, apicibus obtusis vel truncatis, non productis vel subrostratis vel late
rostratis. Pseudosepta curta, c. I:5u longa, planitiei valvari fere parallela. Raphe
filiformis, recta, poris centralibus distantioribus, fissis terminalibus in idem latus
inclinatis. Area axialis angusta ; area centralis parva, circularis vel transverse
elliptica. Noduli terminales minores ; nodulus centralis parva, rectangularis.
Tigilla duo in altero latere inter se angulum c. 30° facientia. Striae leniter radiatae,
in apicibus plerumque parallelae subconvergentesve, 20-25 in 10p, dense punctatae.

jae

Fics. 8-11. Capartogramma crucicula (Grun. ex Cleve) Ross. Fig. 8, valve view (in
B.M. 32046) ; Fig. 9, girdle view (in B.M. 68168) ; Fig. 10, valve view (B.M. 72119,
specim. n. 3) ; Fig. 11, valve view (B.M. 72119, specim. n. 1). (All x 1,000.)

In fresh and slightly brackish waters of temperate and tropical America and of
Africa south of the Sahara, and fossil in pleistocene deposits from the same regions.

U.S.A.: Merrimac River (New Hampshire or Massachusetts) (Coll. P. T. Cleve
n. 729 in Herb. Mus. Stockh., type).

BRAZIL : Amazonas: Rio Purus (7° 35’ S., 65° 22’ W.), from a sandbank,
J. W. H. Trail (B.M. 24530, 24551, 25246, 32043, 32044, 32045, 32046).

TANGANYIKA : Western Province : Lake Tanganyika, Kala Langabo Bay, near
Kigoma, on stems and adventitious roots of Phragmites mauritianus, 30 Jan. 1953,
Ross 1458 (B.M. 72102). Lake Tanganyika, Kigoma, on submerged leaves of

1 This combination is not validly published (see p. 57).

CAPARTOGRAMMA AND SCHIZOSTAURON 61

Paspalidium geminatum, 31 Jan. 1953, Ross 1468 (B.M. 72108, 72109) ; same locality,
on Potamogeton pectinatus, 31 Jan. 1953, Ross 1470 (B.M. 72110, 72111, 72118,
72119, 72120) ; same locality, on Potamogeton schweinfurthti, 31 Jan. 1953, Ross
1472 (B.M. 72127) ; same locality, on Lagarosiphon sp. floating in harbour, 31 Jan.
1953, Ross 1473 (B.M. 72129) ; same locality, on roots of Phragmites mauritianus
c. I m. deep, 31 Jan. 1953, Ross 1474 (B.M. 72131) ; same locality, on leaves c.
120 cm. below surface of water, 31 Jan. 1953, Ross 1475 (B.M. 72133) ; same
locality, on Ceratophyllum demersum in harbour, 31 Jan. 1953, Ross 1477 (B.M.
72137) ; same locality, scrapings from jetty wall, 31 Jan. 1953, Ross 1478 (B.M.
72139) ; same locality, bottom mud in harbour, 2 Feb. 1953, Ross 1512 (B.M. 72141,
72142, 72146) ; same locality, scrapings from hull of M.V. ‘“‘ Sangala”’, 31 Jan.
1953, Ross 1537 (B.M. 72153).

NATAL : Tugela District: On the east of the State Forest between Mtubatuba
and Lake St. Lucia, small brook with nearly still water, 19 July 1957, Cholnoky
TUG. 221 (B.M. 68031). Between the high mountains of Zululand near Eshowe,
little brook, 20 July 1957, Cholnoky TUG. 227 (B.M. 68037). Umlalazi River
near Eshowe, 20 July, 1957, Cholnoky TUG. 228 (B.M. 68038). Near the road fork
leading to Empangeni in the vicinity of the Umhlatuzi River, little brook in rocky
bed, 20 July 1957, Cholnoky TUG. 229 (B.M. 68039). Mpanza brook below the
Greytown-Tugela Ferry road bridge, beds of Cladophora, 21 July 1957, Cholnoky
TUG. 245 (B.M. 68055). Rooispruit Brook between Seven Oaks and Dalton,
26 July 1957, Cholnoky TUG. 281 (B.M. 68081). Between Fown Leas and Rooispruit,
small side brook of the Mbalana River on a rocky bed, 26 July 1957, Cholnoky
TUG. 282 (B.M. 68082). Umlalazi River, at the road bridge between Eshowe and
Melmoth, 27 July 1957, Cholnoky TUG. 285 (B.M. 68085). Nkwaleni Brook
between Melmoth and Eshowe, 27 July 1957, Cholnoky TUG. 286 (B.M. 68086).
Umzimhlanga River between Ixopo and Richmond, 21 July 1958, Cholnoky TUG. 356
(B.M. 68156). Small bridge near Port Shepstone just outside the western side of
the Indian settlement, 21 July 1958, Cholnoky TUG. 361 (B.M. 68161). Amahlongwa
River, at the crossing of the old (westerly) Durban road, 22 July 1958, Cholnoky
TUG. 367 (B.M. 68167). Amanzimtoti River near Illova, 22 July 1958, Cholnoky
TUG. 368 (B.M. 68168). Umgeni River, near the Roberts Construction road bridge
and the trading store, 23 July 1958, Cholnoky TUG. 372 (B.M. 68172). Umgeni
River, by the second road bridge near the Neagle Dam, 23 July 1958, Cholnoky
TUG. 375 (B.M. 68175). Mpanza River, near the Greytown-Tugela Ferry road
bridge, 24 July 1958, Cholnoky TUG. 376 (B.M. 68176). Ample brook, on rock sills
south of Rorke’s Drift, between the Mission near the road fork to Elandskraal,
between Rorke’s Drift and Helpmekaar, and the Drift, 24 July 1958, Cholnoky
TUG. 386 (B.M. 68186). Witfolozi River, near Vryheid, 25 July 1958, Cholnoky
TUG. 390 (B.M. 68190).

FossiL : Pleistocene :

EL SALVADOR : Potonico, freshwater fossil deposit (B.M. 14466 = Temp. & Perag.,
Diatomées, n. 212 ; B.M. 32833 ; B.M. 68366 = Temp. & Perag., Diatomées, ed.
2, nN. 20).

62 CAPARTOGRAMMA AND SCHIZOSTAURON

Most of the previous records of this species have been based on gatherings from
rivers, streams or ponds. I, however, found it frequently in epiphytic and, to a
lesser extent, benthic habitats in Lake Tanganyika at Kigoma. It would seem
therefore to be a species of the layer of diatoms and other algae that grows on higher
plants and other solid substrata in shallow water, and its occasional occurrence in
plankton is fortuitous. In this it resembles C. rhombicum and C. amphoroides,
but these two are confined to Lake Tanganyika, as far as is at present known,
whilst C. crucicula has a much wider geographical range and occurs as frequently
in rivers and streams as in lakes. The records by Zimmermann from Santos,
Brazil, by Frenguelli from the estuary of the River Plate, and by Hustedt from
Venezuela are based on gatherings from estuarine localities, but salinity data are
only available for the last of these ; the water from which this sample was taken
had a sodium chloride content of 1-0 parts per mille. In most of the gatherings
in which I have seen this diatom it has been uncommon.

Cholnoky (Osterr. Bot. Zeitschr. 103 : 87 (1956)) questions the accuracy of
Zanon’s record of this species from Upper Volta, saying : “‘ Zanon... teilt die Art
aus Franzosisch-Mittelafrika (Niger-Fluss) mit, seine Zeichnungen (Il. c., T. 2, F. 12,
zwei Abbildungen) beziehen sich aber auf eine Form aus der Verwandtschaft der
S{tauronets| obtusa Lgst.”” He had, however, failed to notice that there is a misprint
in the text of Zanon’s paper, but not in the explanation of the plate, where t. 2 fig. 12
is stated to represent Stauronets obtusa var. minor and t. 2 fig. 13 Schizostauron
crucicula. The three illustrations which constitute fig. 13 are only about 2 cm.
away from fig. 12, and they are unmistakable representations of C. crucicula.

There is considerable variation in outline in this species, but little in other respects,
except for the occurrence of divided tigilla. The larger specimens (Fig. 8) have
long, broad, rostrate apices, whilst the smaller ones (Figs. 10, 11) are not at all
produced at the apex. There appears to be a continuous variation between the
two extremes. However, the specimens in Cholnoky’s gatherings from the
Tugela District in Natal and mine from Kigoma in Lake Tanganyika appear to
represent very distinct populations. The majority of the Natal specimens are 25
long or more and have long rostrate apices, and the smallest specimens from there
are 18y long and are distinctly subrostrate. The largest of the Lake Tanganyika
specimens is little longer than this and is no more markedly subrostrate, whilst the
smallest ones from there are as little as tou long and have no trace of production
of the apices. Forking or duplication of the tigilla is also frequent in the Lake
Tanganyika specimens (Plate 1 B), but elsewhere it is rare, although it does occur
(Fig. 9). O. Miller found forms similar to the Lake Tanganyika population
in a sample from a pool at Manda (formerly Wiedhafen) on the shores of Lake
Nyasa ; he described them as [Navicula sect.] Schizostauron crucicula forma
obtusa. Somewhat similar though rather larger specimens from Sierra Leone were
named Stauroneis merrimacensis var. intermedia by Woodhead and Tweed. In
view of the identity in appearance of the largest of the Lake Tanganyika specimens
and the smallest of the Natal ones, the present state of our knowledge does not
justify the maintaining of these taxa as distinct.

CAPARTOGRAMMA AND SCHIZOSTAURON 63

As well as the variation with length of valve in the extent to which the apices
are produced, there are also differences in the length-breadth ratio and the convexity
of the valve margin between individuals of the same length. Whilst specimens in
a single gathering usually show considerable uniformity, amongst the different
gatherings a continuous series between the extremes can be found, and therefore
it does not appear justifiable to maintain as distinct [Navicula sect.] Schizostauron
crucicula forma gracilior O. Muell., which is based on slender specimens with almost
straight sides from the plankton of Lake Nyasa and the Songwe River.

Cleve originally intended to use a different epithet for this species and his type
specimen is labelled with this and not Schizostauron crucicula. He gives his reason
for using a different name in a footnote to his protologue which reads : “‘ In printing
this paper I have been informed by Mr. Grunow that he has found the same form
abundantly in pools of the Rio Purus, Brazil, and that he has named it in Linnaean
Society Journ. of May 1880 [.] I consequently adopt his name .”

Until very recently most authors have called this species Stauronetis crucicula, a
later homonym of S. crucicula W. Smith (Synops. Brit. Diat. 1 : 60 (1853)), but
Woodhead and Tweed have recently published a legitimate substitute, S. merri-
macensis. Zimmermann’s independently described Schizostauron brastliense, how-
ever, provides an earlier epithet which would have to be adopted if the species
were placed in the genus Stauroneis. Cholnoky (Hydrobiologia 19 : 104 (1962)),
without referring to the new name proposed by Woodhead and Tweed, recently
suggested that, since the name Stauronets crucicula could not be used for this species,
it should be called S. grunowii Rabenh. (Fl. Eur. Alg. I : 251 (1864)), founded on
S. crucicula var. minuta Grun. (Verh. Zool.-bot. Ges. Wien 10 : 567, t. 6 fig. 15
(1860)) ; contrary to what Cholnoky says, however, this is based on small forms of
Stauroneis crucicula W. Smith (= Navicula crucicula (W. Smith) Donkin) and not of
S. crucicula (Grun. ex Cleve) Comber (= Capartogramma crucicula). More recently
still Cholnoky (Revista Biol. 3 : 69 (1962)) uses the name Stauroneis merrimacensis
for this species without referring to his suggestion that S. grunowz1 is its correct name.

The combination Stauronets crucicula is usually attributed to Cleve, but this is
incorrect, for he did not recognize Stauroneis as a separate genus. In his Synopsis
of the Naviculoid Diatoms his treatment above the specific level is chaotic. His
major groups are all treated taxonomically as if they are of equal status, but nomen-
claturally some are treated as genera and some as sections of Navicula. There is
no description of the genus Navicula as such in the whole work, and the various
sections of that genus described under such headings as Naviculae Fusiformes and
Naviculae Lineolatae are not grouped together but are interspersed among the
accounts of genera he recognized both taxonomically and nomenclaturally. In
the section headed Naviculae Microstigmaticae, with which we are here concerned,
the situation is even more confused. The account begins with a description
of the section, and this is followed by :

“This large section comprises a number of species, hitherto placed in Navicula,
Stauroneis, Pleurostauron, Schizostauron and Schizonema. They may be classed
in the following divisions :

64 CAPARTOGRAMMA AND SCHIZOSTAURON

Stauroneis.

Pleurostauron.

Schizostauron.

Libellus.

. Microstigma.

‘This division of the whole group cannot be completely carried out at present,
as the connecting zones of many species have not been observed. I consequently
class all the forms now in two groups, viz. those with transversely dilated central
nodules in Stawroners, and those without such dilated central nodules in Microstigma ;
at the same time indicating as far as can yet be made out to which of the five groups
named above each species belongs .”

There is no intervening heading between the key to the species and the individual
accounts of them. In these accounts the names are given thus: “S....” or
““S. (Labellus) ...” or “S. (Pleurostauron) ...”. After species n. 27 there is
a heading “‘ Subdivision Schizostauron Grun.’’, under which the species names are
given thus: “S....’’; the six species under this heading include “ S. crucicula’”’.
There is then a further heading : “‘ Subdivisions Microstigma and Libellus ’’, under
which the species names are given as “ N....” or “ N. (Labellus) ...” or “N.
(Schizonema Libellus) ...’’ or, in one case, “‘ N. (Scoliopleura) ...”’. (Elsewhere in
the Synopsis Cleve treats Scoliopleura as a separate genus!) That the “S.” of
“'S. crucicula’’ represents Stauroneis and not Schizostauron is indicated by the
feminine terminations of the adjectival specific epithets under the same heading,
but in the index Schizostauron Cruciculum (sic) is indicated as a correct name and
Stauronets cruciculaasasynonym. Cleve accordingly cannot be held to have regarded
Stauroneis as a genus, and hence the combination Stauronets crucicula is not validly
published in this work.

Boyer (Proc. Acad. Nat. Sci. Philad. 79, Suppl.: 426 (1928)) realized this, but
thought that the combination had first been validly made by himself (Diat. Philad.:
89 (1916)). It was in fact made earlier by Comber, but the Welwitsch specimens to
which he applied it are, as examination of his original preparations shows, Navicula
exiguiformis Hust.

Frenguelli (Rev. Mus. La Plata, Nueva Ser., Bot. 3 : 245 (1941)) suggested that
Pinnularia signata Ehrenb. (Mikrogeologie : t. 34 VI A fig. 7 (1854)) from Florida
might be this species, but this seems unlikely ; Ehrenberg’s figure shows coarse
striae, whilst his optical equipment was such that he would not have been able to
detect those of Capartogramma crucicula, and also the outline of his figure is quite
different from that of this species.

YRwWNH

a”

4. Capartogramma amphoroides Ross, sp. nov. (Plate 1c ; Figs. 12-14.)
Frustulum marginibus longitudinalibus convexis vel arcuatis, cingulo latiore in
parte dorsali quam ventrali, e copulis composito. Valva 20-50u longa, 6-5-9:5u
lata, maxime convexa, lunata, margine ventrali recto vel leniter convexo, aliquando
medio gibboso, apicibus subrostratis. Pseudosepta 2:5—-6:5u longa, planitiei valvari
fere parallela, marginibus interioribus crassis. Raphe filiformis, fere recta, poris

CAPARTOGRAMMA AND SCHIZOSTAURON 65

centralibus distantioribus, fissis terminalibus in idem latus inclinatis. Area axialis
angusta ; area centralis parva, circularis. Noduli terminales minimi ; nodulus
centralis minor, rectangularis. Tigilla duo dorsalia inter se angulum 20°—30°
facientia, duo ventralia multo latius digredentia. Striae leniter radiatae, dorsales
13-22 in 10, ventrales 18-22 in I0u.

Holotype : specimen no. 1! in praep. n. B.M. 72113 in Coll. Diat. Herb. Mus.
Brit.

From the shores of Lake Tanganyika near Kigoma.

—

13

Fics. 12-14. Capartogramma amphoroides Ross. Fig. 12, valve view (holotype) ;
Fig. 13, girdle view (B.M. 72113, specim. n. 5) ; Fig. 14, girdle view (in B.M. 72137).
(All x 1,000.)

TANGANYIKA : Western Province : Lake Tanganyika, Kala Langabo Bay, near
Kigoma, on stems and adventitious roots of Phragmites mauritianus, 30 Jan. 1953,
Ross 1458 (B.M. 72102) ; same locality, on tree root in c. 15 cm. of water, 30 Jan.
1953, Ross 1461 (B.M. 72104). Lake Tanganyika, Kigoma, on submerged leaves of
Paspalidium geminatum, 31 Jan. 1953, Ross 1468 (B.M. 72108, 72109) ; same locality,
on Potamogeton pectinatus, 31 Jan. 1953, Ross 1470 (B.M. 72110, 72111, 72112,
72113) ; same locality, on Potamogeton schweinfurthi, 31 Jan. 1953, Ross 1472
(B.M. 72127) ; same locality, on Lagarosiphon sp. floating in harbour, 31 Jan. 1953,
Ross 1473 (B.M. 72129) ; same locality, on roots of Phragmites mauritianus c. I m.
deep, 31 Jan. 1953, Ross 1474 (B.M. 72131) ; same locality, on leaves c. 120 cm.
below surface of water, 31 Jan. 1953, Ross 1475 (B.M. 72133) ; same locality, mud
from roots c. 120 cm. below surface of water, 31 Jan. 1953, Ross 1476 (B.M. 72135) ;
same locality, on Ceratophyllum demersum in harbour, 31 Jan. 1953, Ross 1477
(B.M. 72137) ; same locality, scrapings from jetty wall, 31 Jan. 1953, Ross 1478
(B.M. 72139) ; same locality, bottom mud in harbour, 2 Feb. 1953, Ross 1512
(B.M. 72141). Lake Tanganyika, c. 1 km. off mouth of Luichi River, on roots
brought up in trawl, 3 Feb. 1953, Ross 1532 (B.M. 72145) ; same locality, mud
from large mussel shell brought up in trawl, 3 Feb. 1953, Ross 1533 (B.M. 72147).
Lake Tanganyika, Kigoma, scrapings from hull of M.V. “ Sangala”’, 31 Jan. 1953,

1 The specimen at the left-hand end of the row as seen under the microscope, the top of the labels
on the slide being to the left.

66 CAPARTOGRAMMA AND SCHIZOSTAURON

Ross 1537 (B.M. 72153). Lake Tanganyika, Katabi Bay, near Kigoma, bottom
sand, 20 m., 4 Feb. 1953, Ross 1538 (B.M. 72155, 72156). Lake Tanganyika,
c. 500 m. offshore between Ujiji and Bangwe Point, on various weeds tangled in
trawl from c. 8 m. deep, 4 Feb. 1953, Ross 1539 (B.M. 72157, 72158) ; same locality,
tangled in meshes of trawl from c. 8 m. deep, 4 Feb. 1953, Ross 1540 (B.M. 72159,
72160). Lake Tanganyika, Bangwe Point, on rocks just below water level, but
exposed during moderate swell, 4 Feb. 1953, Ross 1541 (B.M. 72161). Lake Tan-
ganyika, Kigoma, bottom sand, c. 1:5 m., 4 Feb. 1953, Ross 1549 (B.M. 72171).

This species, like C. rhombicum and C. crucicula, is primarily an inhabitant of the
felt of algae and diatoms that grows in shallow water on solid substrata such as
higher plants, rocks, etc. In my gatherings from Lake Tanganyika it is more
common than any of the other three species of the genus, and it is therefore somewhat
surprising that it has not been found previously either by G. S. West in Cunnington’s
collections or by Kufferath in the material from the Belgian hydrobiological survey
of the lake.

Whilst C. amphoroides differs markedly in symmetry from the other members of
the genus, its resemblance in all other respects is extremely close and there can be
no question but that it is correctly placed here. It is somewhat variable in the
density of striae on the dorsal side of the valve and also in the outline of the frustule,
some specimens being much more inflated than others, but the variation in these
characters is continuous and not of taxonomic significance at any level. The
ventral tigilla of this species are frequently forked (Fig. 14) and occasionally a
dorsal tigillum is forked.

Boyer (Diat. Philad.: 60, t. 16 fig. 13 (1916)) described and figured a specimen
which he identified as the raphe valve of Achnanthes danica (Flogel) Grun. The
description and the outline as shown in the figure are consistent with its being a
raphe valve of A. fimbriata (Grun.) Ross, a species long confused with A. danica
(cf. p. 75 below), but if the central area is correctly drawn it cannot belong to that
species and may well represent an unknown species of Capartogramma with three
struts on either side of the central nodule.

THE IDENTITY OF SCHIZOSTAURON

Schizostauron was established by Grunow (Hedwigia 6 : 28 (1867)) for four marine
species with ovate or broad-lanceolate valves and a central nodule which was
transverse, linear, and bifid at either end. There is a formal latin description of
the genus and of one of the species, S. Jindigianum, the only one found in the material
from Honduras on which he was reporting. The diagnostic characters of the remain-
ing three, S. reichardtianum, S. ovatum and S. fimbriatum, are mentioned in the
discussion in German which follows. Grunow, however, said that he was doubtful
whether it was correct to place S. fimbriatum in the same genus as the other three
species.

Ten years later Grunow published another version of his account of the diatoms
of Honduras. In this he repeated, slightly modified, his latin diagnoses of Schizo-

CAPARTOGRAMMA AND SCHIZOSTAURON 67

stauron and S. lindigianum and gave a latin diagnosis of S. reichardtianum, spelling
the epithets “ Lindigit”’ and “ Reichardtii’”’ respectively. He also figured the two
species (Month. Microsc. Journ. 18 : 181, t. 195, figs. 17, 18 (1877)). Both figures
show the stauros as a narrow transverse band reaching about half-way to the margin
on either side and there forking into curved and tapering bands, whose general
direction is longitudinal.

The types of Grunow’s species are preserved in the Botany Department of the
Naturhistorisches Museum, Vienna, and I have been able to examine both these
and Grunow’s notes and sketches relating to them. On the slide (Coll. Grunow n.
839&) indicated by Grunow as that on which the type of S. lindigianum is to be
found there is only one specimen that agrees with his description and figure. On
close examination, using phase contrast as well as transmitted light, this was found
to be a specimen of Cocconeis dirupta Greg. It consists of a raphe-less valve and
two raphe valves back to back, one of these with its central portion broken away.
There is a transverse fascia on both intact valves, but what Grunow interpreted
as the bifid prolongation of the stauros is no more than a line of contact between
the two valves. This line is obvious because the slide is a dry mount and a certain
amount of dirt has accumulated along it. The expansions of the central area are
also artefacts of a similar nature.

The types of S. reichardtianum and S. ovatum come from a gathering made by
Reichardt in the Adriatic near the town of Mali LoSinj, formerly known as Lussin
Piccolo, on the island of LoSinj in the Gulf of Kvarner. In his second account
of S. reichardtianum Grunow also cited a specimen on Codium bursa from Dalmatia
collected by Dr. Bartsch. Both the type (Coll. Grunow n. 864a) and this latter
specimen (Coll. Grunow n. 1133) are whole frustules of the species of Achnanthes
recently described as new by Proschkina-Lavrenko under the name A. lyrata (Not.
Syst. Sect. Crypt. Inst. Bot. Komarov. 14 : 38, t. 1 figs. 12, 13, t. 2 figs. 18, 19
(r961)). This species is related to A. baldjickui (Brightw.) Grun., A. danica (Fl6égel)
Grun. and A. fimbriata (Grun.) Ross. It has ovate valves with rounded rostrate
apices, striae on the raphe-less valve consisting of single hyaline alveoli and much
finer striae on the raphe valve, whose central area divides on either side into two
curved tapering branches ; between these there is a triangular striate area on which
the striae are not alternately longer and shorter. Grunow clearly did not realize
that he was looking at two valves and he described and figured his specimen as if
it were a single valve with a raphe and the central area of the raphe valve but the
coarser unbroken striae of the raphe-less one. Proschkina-Lavrenko’s interpretation
of the raphe valve seems to be based on a similar error.

Cleve (Ann. & Mag. Nat. Hist., Ser. 7, 10 : 29 (1902)), in the course of a series of
annotations on Karsten’s Die Diatomeen der Kieler Bucht, wrote as follows: “I
have once found in plankton from Skagen a diatom the lower valve of which agreed
with fig. 117, but the upper valve was identical with Grunow’s Schizostauron
Reichardtianum, which thus belongs to Achnanthes, s. 1.” Karsten’s fig. 117 is
labelled Cocconeis apiculata A. Schmidt and depicts a raphe-less and a raphe valve,
both with slightly radiate continuous striae about 10 in rou. It is thus clear that

68 CAPARTOGRAMMA AND SCHIZOSTAURON

the specimen to which Cleve refers was something quite different from Grunow’s
species or was thoroughly misinterpreted by him. Cleve’s remarks do not, of course,
constitute publication of the combination Achnanthes reichardtiana, but this was
validly published by Hustedt (in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 429 (1933)),
who wrongly attributed it to Cleve.

S. ovatum, like S. lindigianum, proves on examination of the type material to be
based on a specimen of Cocconeis dirupta Greg. On this specimen, however, the
marks which Grunow misinterpreted as bifid prolongations of the stauros are inclined
to one another at a much smaller angle and there is no artefact giving the appearance
of expanded axial areas.

Grunow did not designate a type species of Schizostauron, and, since he was
doubtful whether S. fimbriatum was correctly placed in the genus, one of the other
three species originally included in it must be chosen as lectotype. As has been
pointed out, the character which Grunow considered diagnostic of the genus, viz.
the bifid stauros, is possessed by only one of these, S. reichardtianum, and this must
accordingly be chosen. In the other two species the bifid stauros is only simulated
by an artefact and neither of them can be accepted as the type of the genus, even
although they have never been removed from Schizostauron whilst S. reichardtianum
has been transferred to Achnanthes. Since the correct taxonomic position of the
type species is in that genus, Schizostauron must be regarded as a synonym of
Achnanthes.

A number of species other than the three considered above have been referred
to Schizostauron by various authors. Whilst two are, like the type of the genus,
species of Achnanthes, most belong to other genera.

S. fimbriatum Grun. (Hedwigia 6 : 28 (1867)), doubtfully referred to the genus
Schizostauron when it was first established, proves on examination of the type
material (Coll. Grunow n. 869, Porto Zubzamki, SuSak Island, Gulf of Kvarner,
Jugoslavia, leg. Reichardt) to be the raphe valve of the species currently known as
Achnanthes manifera Brun. Grunow’s name provides the earliest specific epithet
for this, and the necessary new combination is accordingly made below (p. 72).
This species has been confused with A. danica (Flégel) Grun., which has in its turn
been confused with the diatom hitherto known as A. lorenziana (Grun.) Cleve but
named here (p. 82) A. baldjickii subsp. lorenziana (Grun.) Ross. An account of
the confusion between these species accompanies their synonymies and descriptions
(pp. 72-84 below).

Cleve, in his original account of S. andicola (Ofvers. K. Vetensk.-Akad. Férhandl.
Stockh. 38 (10) : 12, t. 16 fig. 8 (1882)), expressed doubt as to whether it was
correctly placed in Schizostauron. It was subsequently transferred to Achnanthes
by Ostrup (Danske Diat.: 128 (1910)), who referred it to A. hungarica (Grun.)
Grun. as a distinct variety. In the following year Hustedt (Abhandl. Naturwiss.
Ver. Bremen 20 : 279 (1911)) treated it as a distinct species of Achnanthes. Exami-
nation of the type material shows that the species was based on large specimens of
A. hungarica.

De Toni (Nuova Notarisia [1] : 196 (18g0)) transferred Stawroneis tatrica Gutw. to

CAPARTOGRAMMA AND SCHIZOSTAURON 69

Schizostauron. Cleve, in his Synopsis of the Naviculoid Diatoms, treated this as a
synonym of Navicula pupula Kiitz., and it would appear from Gutwinsky’s account
and figure (Sprawozd. Kom. Fizyjogr. 25 : (24), t. 1 fig. 20 (1890)) that he was
correct in so doing.

De Toni (Syll. Alg. 2 : 225 (1891)) also transferred Stauroneis verbania De Not.
to Schizostauron. Examination of the type material (Erb. Critt. Ital., Ser. 2:
n. 434 (1871)) shows that this too is Navicula pupula.

Stauronets sagitta Cleve (K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 18 (5) :
15, t. 3 fig. 45 (1881)) is listed by Cleve in his Synopszs (op. cit. 26 (2) : 151 (1894))
under the subdivision Schizostauron of the Naviculae Microstigmaticae. Hustedt
(in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 811 (1959)) reduces this taxon to a variety
of Stauroneis smith Grun. As explained above (p. 53), that species resembles
Capartogramma more closely than it does the type species of Stawroneis, but there is
not yet sufficient information to justify its transfer either to Capartogramma or to a
new genus.

When the original material of Schizostauron rhombicum M. Perag. (in Temp. &
Perag., Diatomées, ed. 2 : 398, n. 825 (1913)) was examined, it was found to be a
Navicula with a transverse fascia widening outwards but not reaching the margin.
This species has apparently not been reported by any other author and, as the name
Navicula rhombica is pre-occupied (Gregory in Quart. Journ. Microsc. Sci. 3 : 40
(1855)), a new name is proposed for it on p. g1 below.

The only other species that has been referred to Schizostauron is S. portaregtense
A. Berg (Ark. Bot., Andra Ser., 2 : 30, 36, t. 4 fig. 57 a-c (1952)). I cannot find
anything to match Berg’s illustration in his type material. He reports, however,
that he examined some uncleaned material mounted in Viscol of the gatherings
in which he found this species, and I believe that his name is based on specimens of
Navicula rhyncocephala var. amphiceros (Kiitz.) Grun. with remains of cell contents
simulating a stauros and apical septa. The identity of his species must, however,
remain doubtful, although it is clearly neither a Capartogramma nor a Stauronets.

Species Investigated—Synonymies, Descriptions and Notes

CocconEIS DIRUPTA Greg. in Trans. R. Soc. Edin. 21 : 491, t. 9 fig. 25 (1857).—Hust.
in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 354, fig. 809 (1933).

Cocconeis diaphana var. 8 W. Smith, Synops. Brit. Diat. 1 : 22, t. 33 fig. 254 6 (1853).

Cocconeis flexella Janisch & Rabenh. in Rabenh., Beitr. Kenntn. Verbr. Alg. 1 : 7, t. 1
fig. 11 (1863).

Schizostauron lindigianum Grun. in Hedwigia 6 : 28 (1867) ; in Month. Microsc. Journ.
18 : 181, t. 195 fig. 17 (1877).

Schizostauron ovatum Grun. in Hedwigia 6 : 28 (1867).

Cocconeis beltmeyeri Janisch ex A. Schmidt, Atl. Diat.: t. 196 fig. 23 (1894).

Cocconeis delicata A. Schmidt, tab. cit. fig. 24 (1894).

Stauroneis lindigiana (Grun.) Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Fdljd,
26 (2) : 151 (1894).

Stauroneis ovata (Grun.) Cleve, loc. cit. (1894).1

Stauroneis lindigiana (Grun.) Mills, Index Gen. & Sp. Diat.: 1462 (1934).

Stauroneis ovata (Grun.) Mills, op. cit.: 1464 (1934).

1 These combinations, although used here by Cleve, were not validly published by him (see pp. 63-64).

70 CAPARTOGRAMMA AND SCHIZOSTAURON

For description and figure, see Hustedt (loc. cit.).

A cosmopolitan marine littoral epiphyte.

ENGLAND : Sidmouth, Devon, Miss Cutler (B.M. 19589, C. diaphana var. 8
W. Smith).

SCOTLAND : Corriegills, Isle of Arran, on Corallina, Miles (B.M. 1420, isotype).

JuGosLAvIA : Mali LoSinj, LoSinj Island, Gulf of Kvarner, Reichardt (Coll.
Grunow n. 864b in Herb. Vienna, holotype of Schizostauron ovatum).

Honpuras:: On Sargassum, Lindig (Coll. Grunow n. 839% in Herb. Vienna,
holotype of Schizostauron lindigianum).

W. Smith, in his protologue of Cocconeis diaphana, cites two gatherings, one from
Jersey and one from Sidmouth, without indicating which represents his var. 8.
However, the material from Jersey matches his fig. 254, and that from Sidmouth
his fig. 2548 and his description of var. 8, which is said to have a stauros. The
Jersey specimens are identical with the type material of C. molesta Kiitz., showing
that C. diaphana is correctly regarded as a synonym of that species, whilst those
from Sidmouth belong to the species later described by Gregory as C. dirupta.

ACHNANTHES REICHARDTIANA (Grun.) Hust. in Rabenh., Krypt.-Fl. Deutsch. 7 (2) :
429 (1933).—Mills, Index Gen. & Sp. Diat.: 1439 (1934), 1699 (1935). (Figs.
15, 16.)

Schizostauron veichardtianum Grun. in Hedwigia 6 : 28 (1867) ; in Month. Microsc. Journ.
18 : 181, t. 195 fig. 18 (1877).

Stauroneis veichardtiana (Grun.) Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Foljd,
26 (2) : 151 (1894).3

Stauroneis veichardtiana (Grun.) Mills, op. cit.: 1468 (1934).

Achnanthes lyvata Proschk.-Lavr. in Not. Syst. Sect. Crypt. Inst. Bot. Komarov. 14: 38,
t,t figs. 12.13, t) 2 figs.-18, 10) (1961).

—

Fe
ie

Sup "

fon)

Fics. 15-16. Achnanthes reichardtiana (Grun.) Hust. (holotype). Fig. 15, raphe valve ;
Fig. 16, raphe-less valve. (Both x 1,000.)

—_

Valve broadly elliptic with short subrostrate to rostrate apices, 30-40 long,
18-23 broad. Raphe-less valve with a linear or very narrowly lanceolate pseudo-
raphe and a narrow central area tapering outwards ; striae slightly radiate, slightly
curved, 8-11 in 10y, each consisting of a single hyaline alveolus, rather distant
throughout most of their length but widening and becoming almost contiguous at
the margin. Raphe valve with a straight filiform raphe lying in a thickened rib ;

1 This combination, although used here by Cleve, was not validly published by him (see pp. 63-64).

CAPARTOGRAMMA AND SCHIZOSTAURON 71

central pores approximate ; terminal nodules small ; axial area linear, narrow ;
central area a narrow transverse fascia bifurcating half-way to the margin on either
side into two curved tapering branches ; striae punctate, strongly radiate through-
out except within the bifurcation of the central area, sinuous adjacent to the central
area, curved near the apices, 22-24 in 10, puncta about as distant as the striae.

Marine, from the coast of Jugoslavia and from the Black Sea.

JUGOSLAVIA : Mali LoSinj, LoSinj Island, Gulf of Kvarner, Reichardt (Coll.
Grunow n. 864a in Herb. Vienna, holotype). Dalmatia, on Codium bursa, Bartsch
(Coll. Grunow n. 1133 in Herb. Vienna).

This is an uncommon species, only a single example being present in each of the
two preparations on which it occurs, and in both cases the specimen is somewhat
broken. In the material from Mali LoSinj the related species Achnanthes fimbriata
and A. baldjickit also occur and are more common. The only other record is
Proschkina-Lavrenko’s from the island of Tendra in the Black Sea, U.S.S.R.

A. reichardttana has a raphe-less valve closely resembling that of A. baldjickit,
the striae each consisting of a single alveolus without secondary markings visible
under the light microscope ; Proschkina-Lavrenko comments on this similarity.
The raphe valve, however, is much more like that of A. fimbriata, the striae being
close and not double near the margin nor alternating there with costae. It was
not easy to be certain of some details of the structure of the raphe valve ; the only
specimens available are whole frustules mounted in air, and the coarser structure
of the raphe-less valve obscures the finer markings of the raphe valve. In conse-
quence it was impossible to see whether the marginal alveoli of the raphe valve
were elongated, as they are in A. fimbriata and A. danica.

According to Proschkina-Lavrenko, the striae on the raphe valve are considerably
more widely spaced than I found in the Adriatic specimens. Moreover, her drawing
(t. x fig. 13) of this valve shows the striae as only slightly radiate and interrupted
by the branches of the central area. However, her photograph that purports to
be of the raphe valve (t. 2 fig. 19), like Grunow’s original illustration of the species,
shows the raphe and central area superimposed on the markings of the raphe-less
valve, which appear interrupted as a result of interference from the much finer
striae of the raphe valve. Proschkina-Lavrenko, like Grunow, seems not to have
detected these.

The combination A. reichardtiana is usually attributed to Cleve (Ann. & Mag.
Nat. Hist., Ser. 7, 10 : 29 (1902)), but, as is pointed out above (p. 68), he did not
publish it ; this was first done by Hustedt. Cleve did publish the combination
Stauronets reichardtiana, but not validly (cf. pp. 63-64 above). Mills was apparently
the first to give the combination valid publication, but he appears to have done so
by oversight. In the entry for Schizostauron reichardtianum on pp. 1438-1439 of
his Index he indicates that Achnanthes reichardtiana is the correct name for the
species and does not cite Stauroneis reichardtiana as a synonym. On p. 1468 he
lists Stauroneis reichardtiana as a correct name with Schizostauron reichardtianum
as a synonym but no mention of Achnanthes reichardtiana. In the addenda, on
p. 1699, he lists A. reichardtiana as a correct name, with Schizostauron reichardiianum

72 CAPARTOGRAMMA AND SCHIZOSTAURON

as the only synonym. Hustedt’s treatment of Achnanthes in the Kryptogamen-
Flora was published during the period when Mills’s Index was coming out, and it
would seem that Mills accepted Cleve’s and Hustedt’s view that the species belonged
to that genus but failed to correct the entry under Stauroneis that referred to it.

Achnanthes fimbriata (Grun.) Ross, comb. nov. (Figs. 17-22.)

Schizostauron fimbriatum Grun. in Hedwigia 6 : 28 (1867).

Achnanthes heteropsis Grun. ex Cleve & Moller, Diatoms 3 : 6, ns. 154, 155 (1878), nom.
nud.

Navicula fimbriata Grun. ex Cleve & Moller, loc. cit. (1878), nom. syn.

Stauronets cornuta Leud.-Fortm., Cat. Diat. Ceyl.! : 37, t. 3 fig. 36 (1879).

Achnanthes danica Grun. in K. Svensk. Vetensk.-Akad. Handl., Ny Féljd, 17 (2) : 21 (1880)
pro parte ; in Van Heurck, Typ. Synops. Diat. Belg.: 121, n. 545 (1887) ; non Cocconeis
danica Flogel.—Cleve & Moller, Diatoms 6: 3, n. 311, 4, n. 320 (1882).—Cleve in
K. Svensk. Vetensk.-Akad. Handl., Ny Féljd, 27 (3) : 186 (1896)—H. & M. Perag.,
Diat. Mar. France : 7, t. 2 fig. 3 (1897) pro parte.—Temp. & Perag., Diatomées, ed. 2 :
185, ns. 346-349 (1909).

Navicula vaszaryi Pant., Beitr. Kenntn. Foss. Bacill. Ung. 3 : t. 16 fig. 239 (1892) ; Beschr.
Neu. Bacill. Pars III Beitr. Kenntn. Foss. Bacill. Ung.: 83 (1905).

Achnanthes vaszaryi (Pant.) Cleve in Le Diatomiste 2 : 57 (1893) excl. var. ovegonica
Cleve ; in K. Svensk. Vetensk.-Akad. Handl., Ny Féljd, 27 (3) : 183 (1896).

Achnanthes danica var. maxima Cleve ex A. Schmidt, Atl. Diat.: t. 198 figs. 60, 61 (1895).—
Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 27 (3) : 186, t. 3 figs. 8, 9 (1896)

““ major ’’).

Pee eas Brun in Le Diatomiste 2 : t. 16 figs. 86, 87 (1895).—Boyer in Proc.
Acad. Nat. Sci. Philad. 79, Suppl.: 235 (1927).—Freng. in Chiov., Fl. Somala : 394, t. 49
fig. 14 (1929).—Hust. in A. Schmidt, Atl. Diat.: t. 415 figs. 9-18 (1937) ; in Bull. Duke
Univ. Mar. Sta. 6 : 18, t. 6 figs. 1-8 (1955).

Achnanthes stroemii Hust. in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 393, fig. 841B (1933)
(‘ Strémi ”’).

Valves ovate to rhombic-lanceolate, subrostrate, 20-120n long, 7—40y broad,
often with one margin slightly gibbous opposite the central nodule. Raphe-less
valve with the pseudoraphe linear or slightly widened in the central part of the valve
and with a small hyaline space at each apex ; striae distant, slightly to moderately
radiate, sometimes a few opposite the central nodule short, 54-15 in roy, alveolate ;
alveoli several per stria, those adjacent to the pseudoraphe always considerably
longer than broad and usually the longest in the stria ; divisions between the alveoli
transverse or occasionally oblique but never bifurcating or anastomosing ; faint
continuations of the striae present on the central portion of the pseudoraphe on
those specimens where this is widened. Raphe valve with a straight filiform
raphe with approximate central pores ; terminal nodules small, terminal fissures
hooked ; axial area moderately broad at the centre, tapering uniformly to the
apices ; central area transverse, expanded outwards or slightly narrowed outwards
and then expanded, a half of the width of the valve or more, its outer margin ragged ;
striae arcuate, strongly radiate throughout, especially near the apices, longer and

1 This publication is a reprint from Mémoires de la Société d’Emulation des Cétes-du-Nord. I have
been unable to trace a copy of the original.

CAPARTOGRAMMA AND SCHIZOSTAURON 73

shorter opposite the central nodule, 12-30 in roy in the central part of the valve,
closer towards the apices, alveolate ; alveoli punctiform and scarcely longer than
broad except for the marginal one of each stria, which is 1—4y long.

From the coasts of the North Atlantic, the Mediterranean and the Indian Ocean
and the western border of the Pacific Ocean, and fossil from marine and brackish
deposits in eastern Europe.

me DOS SSe Slee

—
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S
SS
18
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SSocn ———
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boOOo — toler =
ene goon SS
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e7 19

Fics. 17-22. Achnanthes fimbriata (Grun.) Ross. Fig. 17, raphe valve (in B.M. 13079)
(x 1,000) ; Fig. 18, detail of same valve as Fig. 17 (x 2,000) ; Fig. 19, raphe-less
valve (in B.M. 13079) (x 1,000) ; Fig. 20, raphe valve (B.M. 37339, specim. n. 4)
(x 1,000) ; Fig. 21, detail of same valve as Fig. 20 (x 2,000) ; Fig. 22, raphe-less
valve (Coll. F. W. Adams n. Bess. 896, specim. n. 1) ( x 1,000).

JuGosLAvIA : Porto Zubzamki, SuSak Island, Gulf of Kvarner, Reichardt (Coll.
Grunow ns. 869, 869b in Herb. Vienna, syntypes). Mali LoSinj, LoSinj Island,
Gulf of Kvarner, Reichardt (Coll. Grunow ns. 864a, 864b in Herb. Vienna).

SPAIN : Balearic Islands (B.M. 36256 = Cleve & Moller, Diatoms, n. 154).

ITALy : Capri (B.M. 33068).

Norway : Grip (Cleve & Moller, Diatoms, n. 311 in Coll. F. W. Adams in Herb.
Brit. Mus.).

U.S.A.: Connecticut : Morris Cove (B.M. 14577 = Temp. & Perag., Diatomées,

74 CAPARTOGRAMMA AND SCHIZOSTAURON

n. 323 ; B.M.69301 = Temp. & Perag., Diatomées, ed. 2, n. 949 ; isotypes of
A. mantfera). Florida : Pensacola (B.M. 13079 = Cleve & Moller, Diatoms, n. 320).

ADEN : Harbour mud, Vozgé (Coll. Voigt n. 34099).

CEYLON : Colombo, Craven (B.M. 8986, syntype of Stauronets cornuta) ; same
locality, harbour mud, Voigt (Coll. Voigt ns. 14099, 30056).

INDONESIA : Celebes, from holothurians (Coll. Voigt n. 29018).

CuiInA : Kwangtung : Hainan, from holothurians (Coll. Voigt ns. 36053, 36060).
Chekiang : Chusan Archipelago, Voigt (Coll. Voigt n. 3040).

JAPAN : Kyushu : Nagasaki, off shells, Vozgt (Coll. Voigt n. 25044).

AUSTRALIA : Queensland : Flinder’s Passage (B.M. 31942).

Fossil : Miocene :

HunGARY : Nyermegy (B.M. 37338 ; B.M. 68520, 68521 = Temp. & Perag.,

Diatomées, ed. 2, ns. 174, 175).
Pliocene :

BuLGARIA : Balchik (B.M. 2314; B.M. 26856, 26857 = Van Heurck, Typ.
Synops. Diat. Belg., ns. 545, 546 ; B.M. 30827, 31384, 32629, 32630, 32633, 32836,
37339, 42089 ; Coll. F. W. Adams ns. A 167, Bess. 895, Bess. 896 in Herb. Brit.
Mus.).

HUNGARY: Bremia, near Kavna (B.M. 14757, 14758 = Temp. & Perag.,
Diatomées, ns. 503, 504; B.M. 33302, 33303; B.M. 68694 = Temp. & Perag.,
Diatomeées, ed. 2, n. 348). Karand (B.M. 70893).

As will be seen, the range of density of striation on both the raphe-less and the
raphe valves of this species is very wide. It is found to vary continuously, however,
when a considerable number of specimens from a variety of localities is examined,
and the variation on the two valves is not closely correlated. There are nevertheless
some differences between the ranges found in recent and fossil specimens, although
there is much overlap. Recent specimens (Figs. 17-19) range in size from 20u
by 7u to 75u by 26yu, whilst the fossil ones (Figs. 20-22) range from 24u by IIp
to 120%. by 40 ; the striae on the raphe-less valve of recent specimens range
from 5°5 to 14 in 10y, whilst those on fossil ones range only from 7:5 to 13 in Ioy ;
on raphe valves the striae of recent specimens range from 18 to 30 in Io in the centre
part of the valve whilst those of fossil specimens range from 12 to 21in rou. Another
variable feature is the direction of the striae on the raphe-less valve ; in some speci-
mens these are definitely although not strongly radiate and a few short striae are
present in the centre of the valve, whilst others have almost parallel striae and no
short ones. Short striae are present on all fossil specimens but only on some recent
ones. These differences between fossil and recent specimens are too indefinite to
serve as a basis for an infraspecific division, and accordingly Achnanthes danica
var. maxima, proposed by Cleve for fossil specimens of this species from Balchik,
is not maintained here as a separate variety.

The striae on the raphe-less valve of this species are always separated by a distinct
gap, usually equal to at least half their own width, but in the coarsest specimens
occasionally somewhat less. On most of Hustedt’s figures of this valve of the species
(viz.: A. Schmidt, Atl. Diat.: t. 415 figs. 9, 11, 13-15, 18 (1937).; Bull. Duke Univ.

CAPARTOGRAMMA AND SCHIZOSTAURON 75

Mar. Sta. 6: t. 6 figs. 5, 7, 8 (1955)) they are represented as contiguous. Whilst
this may to some extent be due to the drawing convention adopted, it is misleading.

Achnanthes fimbriata is less common in the miocene and pliocene deposits from
Hungary than it is in the pliocene deposit from Balchik, Bulgaria, and even there
it is infrequent. It is not common, either, in any of the recent gatherings in which
I have seen it, but nevertheless it has a wide distribution, from the Atlantic Coast
of North America, through Europe and the Mediterranean to Ceylon, the East
Indies and the western borders of the Pacific.

Grunow expressed doubt about the correctness of referring this species to
Schizostauron when first describing it, and by 1877 he had decided that it did not
belong there ; the phrase in the original diagnosis of the genus which especially
refers to it, “ (vel laciniato fimbriato?) ”’, was deleted from the version published
in that year (Month. Microsc. Journ. 18 : 181 (1877)). Grunow’s notes show that
when he first decided that the species was not a Schizostauron he considered that it
should be placed in Navicula but that he then found whole frustules in the material
from the Balearic Islands issued as Cleve & Moller, Diatoms, ns. 154, 155 (1878),
and realized that it was an Achnanthes. He apparently considered “‘ fimbriata ”’
an inappropriate epithet for the species when it was placed in this genus. The list
of species in the preparations forming the third series of Cleve and Mdller’s Diatoms,
which includes ns. 154, 155, is headed “ All the slides have been examined by M.
Grunow ’’, and the list for those numbers includes ‘‘ Achnanthes heteropsis Grun.
(= Navicula fimbriata Grun.)’”’. Neither of these names had been published pre-
viously and both must accordingly be regarded as nomina nuda ; the identity of
epithet is insufficient to connect them with the previously published description of
Schizostauron fimbriatum Grun., although Grunow’s manuscript notes show that
they were new names for that species.

Between 1878 and 1880 Grunow formed the opinion that this species was identical
with Cocconets danica Flogel and he accordingly made the new combination Ach-
nanthes danica, quoting A. heteropsis as a synonym. This mistake was probably
the result of his receiving, apparently from Weissflog, material of a further gathering
from the Balearic Islands in which there was true A. danica but no A. fimbriata.
From his notes it appears that he did not at any stage associate the raphe valves
he called Schizostauron fimbriatum in the material from the Gulf of Kvarner with the
raphe-less valves also present in his preparations of these gatherings. When he
found specimens in the Cleve and Moller material from the Balearic Islands that
caused him to decide that his S. fimbriatum was the raphe valve of an Achnanthes,
he made no drawing of the raphe-less valve. He did, however, make drawings of
both valves of A. danica from specimens in the Weissflog material and, misled by
the similarity of the raphe valves, failed to realize that he was dealing with a
different species from that in Cleve and Moller’s material from the same locality
and also in their material from Pensacola, Florida, which he saw at about the same
time. Asa result of Grunow’s error, A. fimbriata and A. danica have been confused
ever since 1880. Thus Cleve took specimens of A. fimbriata in the material from the
Balearic Islands issued as Cleve & Moller, Diatoms, ns. 154, 155, as representative

76 CAPARTOGRAMMA AND SCHIZOSTAURON

of A. danica, and consequently in the Synopsis of the Naviculoid Diatoms he applied
that name to A. fimbriata and misidentified true A. danica with A. lorenziana
(Grun.) Cleve (= A. baldjickit subsp. lorenziana (Grun.) Ross). The way in which
the name A. danica has been applied is analysed in the account of that species below
(pp. 79-81).

When Brun found specimens of this species in Tempére’s material from Morris
Cove, Connecticut, he realized that they were not conspecific with Cocconets danica
and so described them as a separate species, Achnanthes manifera. Such other
authors as have recognized that this species is distinct from A. danica have, in general,
used Brun’s name for it, but European specimens have for the most part continued
to be identified as A. danica. The exceptions are those from Norway described
by Hustedt as the separate species A. stroemi1, and the two from the Balearic
Islands figured by him (in A. Schmidt, Atl. Diat.: t. 415 figs. 15, 16 (1937)) as
A. manifera.

Cleve (in Nordensk., Vega-Exped. Vet. Iaktt. 3 : 512 (1883)) was the first to treat
Stauronets cornuta Leud.-Fortm. as a synonym of A. danica. Leuduger-Fortmorel’s
species is based on a raphe valve and both A. danica and A. fimbriata occur in the
material from Ceylon on which he was reporting. It is therefore impossible to be
completely certain to which of the two species his type belongs. Nevertheless, the
central area shown in his figure strongly suggests A. fimbriata rather than A. danica
and his inability to resolve the striae points in the same direction.

Navicula vaszaryt Pant. from the Hungarian miocene is clearly based on a raphe
valve of Achnanthes fimbriata, which occurs in the diatomite from Nyermegy from
which Pantocsek’s type came. Cleve transferred the species to Achnanthes on the
basis of the resemblance of the raphe valve to that of an Achnanthes from Oregon
which has a very different raphe-less valve and is clearly unrelated.

Frenguelli (in Chiov., Fl. Somala : 394, t. 49 fig. 14 (1929)) describes and figures a
raphe valve from near Mogadiscio in Somalia under the name Achnanthes mantfera.
As Hustedt (Bull. Duke Univ. Mar. Sta. 6 : 18 (1955)) has pointed out, this cannot be
identified with absolute certainty in the absence of a raphe-less valve, but its central
gibbosity, the shape of its central area, and the closeness of its striae together make
it highly likely that it is A. fimbriata and not A. danica. MHustedt’s (loc. cit.)
statement that according to Frenguelli the striae on his specimen are I0-II in Iou
is an error ; Frenguelli says that this is the density of the striae on the raphe valve
of A. baldjickiit (Brightw.) Grun., with which he compares A. manifera, and that
those on his own specimen are 22 in I0uw.

Achnanthes stroemii Hust. is founded on specimens of this species with rather
close striae on the raphe-less valve and not very close ones on the raphe valve.
His type came from Nordasot, Norway. Specimens closely resembling Hustedt’s
description and figure occur in Cleve & Moller, Diatoms, n. 311, from Grip, Norway,
which is listed as containing A. danica.

ACHNANTHES DANICA (Flégel) Grun. in K. Svensk. Vetensk.-Akad. Handl., Ny
Foéljd, 17 (2) : 21 (1880) excl. syn. Achnanthes heteropsis Grun. et specim. ex
Mediterraneo (partim), Adria et Florida.—A. Schmidt, Atl. Diat.: t. 198 fig. 50

CAPARTOGRAMMA AND SCHIZOSTAURON 77

(1895).—H. & M. Perag., Diat. Mar. France : 7, t. 2 figs. 1, 2 (1897) excl. t. 2
fig. 3.—Hust. in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 396, fig. 845 (1933) excl.
syn. Schizostauron fimbriatum Grun., Achnanthes heteropsis Grun., Stauroneis
cornuta Leud.-Fortm. et specim. ex Cleve & Mller, Diatoms, et Temp. & Perag.,
Diatomées, ed. 2.—Cleve-Euler in K. Svensk. Vetensk.-Akad. Handl., Ser. 4, 4
(5) : 18, fig. 510 (1953).—Job, Suppl. 1 List Diat. Alexandria : [4] (1961). (Figs.
23-26.)

Cocconeis danica Flégel in Jahresber. Comm. Wiss. Unters. Deutsch. Meere Kiel 1 : 91,
fig. 14 (1873).—Van Heurck, Treat. Diat., transl. Baxter : 289 (1896) excl. fig.; in
Microgr. Prep. 13 : 83 (1905).

Achnanthes lorenziana Grun. ex A. Schmidt, Atl. Diat.: t. 198 fig. 59 (1895).

Achnanthes lorenziana Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 27 (3) : 186
(1896) pro parte ; non Rhaphoneis lorenziana Grun.—Hust., tom. cit.: 395 (1933) quoad
descr. valvae cum raphide.—Cleve-Euler, loc. cit. (1953) quoad descr. valvae cum
raphide.—Job, List Diat. Alexandria : [3] (1954).

23

Fics. 23-26. Achnanthes danica (Flégel) Grun. Fig. 23, raphe valve (in B.M. 71125)
(x 1,000) ; Fig. 24, detail of same valve as Fig. 23 (x 2,000) ; Fig. 25, raphe-less
valve (in B.M. 71125) (x 1,000) ; Fig. 26, detail of same valve as Fig. 25 (x 2,000).

Valves rhombic-lanceolate, subrostrate, 50-65u long, 20-25u broad. Raphe-less
valve with the pseudoraphe very narrowly lanceolate and a small hyaline space at
each apex ; striae distant, parallel or very slightly radiate, 4-5-6 in 10u, each con-
sisting of a single alveolus with oblique or transverse, sometimes bifurcating or
anastomosing, secondary costae about 18-20 in Iou. Raphe valve with a straight
filiform raphe with approximate central pores ; terminal nodules small, terminal
fissures hooked ; axial area narrow, tapering uniformly to the apices ; central area
transverse, rhombic, indefinitely bounded at its outer angles ; striae strongly
radiate, longer and shorter in the centre of the valve, 16-20 in I0y, somewhat
closer towards the apices ; alveoli mostly punctiform and 13-17 in Ioy, but those
adjacent to the margin 1-5-2u long ; striae near the apices consisting of a single
alveolus.

From the coasts of western Europe, the Mediterranean and Ceylon, and fossil
from southern Sweden (fide Cleve-Euler ; a doubtful record, cf. p. 81).

78 CAPARTOGRAMMA AND SCHIZOSTAURON

ITaLy : Capri (B.M. 33068, 33069, 33426, 71121, 71122, 71123, 71124, 71125).

SPAIN : Balearic Islands (B.M. 57563, coll. F. W. Adams ns. E 230, E 231 in Herb.
Brit. Mus.).

Ecypt : Ramleh, near Alexandria, 24 Sept. 1952, H. S. Job (Coll. H. S. Job
n. 438) ; same locality, 27 Nov. 1952, H. S. Job (Coll. H. S. Job n. 446) ; same
locality, 5 Apr. 1953, H. S. Job (Coll. H. S. Job n. 468). Rushdy, near Alexandria,
19 Nov. 1958, H. S. Job (Coll. H. S. Job n. 253).

CEYLON : Colombo, Craven (B.M. 8986).

This species, although widespread, is far from common, a fact which probably
accounts for the inaccuracies in the accounts of it in the literature and for the long
confusion of it with Achnanthes fimbriata (Grun.) Ross. When the raphe-less valve
is observed under the microscope with a lens of less aperture than N.A. 1-3, or when
such a lens is used with an inadequate illuminating cone, the striae appear to have
a double row of puncta, in places alternating, in others opposite. When a lens of
N.A. 1°3 or more is used with an adequate illuminating cone, however, it becomes
apparent that this appearance is an optical artefact and that the true structure
is as shown in Fig. 26. Similarly, careful observation with a lens of adequate
aperture shows that the striae of the raphe valve are not regularly punctate through-
out, as depicted by Hustedt, but that there is a row of long alveoli at the margin
(Fig. 24). |

Although the raphe valve is very similar to that of A. fimbriata, the central area
is smaller and does not definitely expand outward, as that of A. fimbriata always
does.

The way in which Grunow confused this species and A. fimbriata has been dis-
cussed under that species (p. 75 above). It has also been confused with A. baldjickit
subsp. lorenziana (Grun.) Ross (A. lorenziana (Grun.) Cleve), firstly by Pantocsek,
who misidentified specimens of that subspecies as A. danica, then by Cleve, who
did the reverse. His account of A. lorenziana in the Synopsis of the Naviculoid
Diatoms is based almost entirely on A. danica. Thus he says of the raphe-less valve
“‘ Intercostal spaces finely punctate (puncta about 23 in 0,or mm.), frequently appar-
ently smooth ’’, and his description of the raphe valve would apply to that of A.
danica or A. fimbriata. The localities he cites for the species are ‘‘ Barcelona, Balearic
Islands! Adriatic (Grun.), Bab el Mandeb! Macassar Straits (Grove Coll.)! ’’ and he
adds the remark : ‘‘ I have seen an entire frustule in Grove’s collection, but the
coarse structure of the upper valve made the examination of the lower very difficult.”
No specimen to which this remark could refer is to be found now in Grove’s collection.
About 1890 Thum issued slides of material from the Balearic Islands containing
both A. danica and A. baldjickii subsp. lorenziana, but no A. fimbriata, and Cleve
is certain to have seen some of these. As has already been pointed out (p. 76 above),
he applied the name A. danica to A. fimbriata and, at least as far as the Balearic
Island material was concerned, he seems to have identified as A. lorenziana the raphe-
less valves of both A. baldjickii subsp. lorenziana, in which the outer membrane to
the single alveolus forming each stria is smooth, and A. danica, in which that mem-
brane has anastomosing secondary thickening. With these raphe-less valves he

CAPARTOGRAMMA AND SCHIZOSTAURON 79

associated the raphe valves of A. danica, and he apparently never saw a raphe valve
of A. baldjickwt subsp. lorenziana in this material.

Before part II of the Synopsis of the Naviculoid Diatoms appeared, but during its
preparation, plate 198 of A. Schmidt’s Atlas was published. Figure 59 on this plate
is undoubtedly of a raphe-less valve of A. danica ; its legend is “‘ Nordsee, nach
Cleve A[chnanthes| Lorenziana Grunow’’. As the name Achnanthes lorenziana
had not by then been published, and as the figure is sufficient to validate its publica-
tion there, Achnanthes lorenziana Grun. ex A. Schmidt (Atl. Diat.: t. 198 fig. 59
(1895)) is a legitimate name, typified by the North Sea specimen figured by Schmidt,
a synonym of A. danica (Flogel) Grun., and an earlier homonym of A. lorenziana
(Grun.) Cleve (K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 27 (3) : 186 (1896)),
whose basionym is Rhaphonets lorenziana Grun.

In view of the way in which the name Achnanthes danica has been so frequently
misapplied, it will probably be useful to present here an analysis of all the accounts
and records which have been published under it :

Grunow in K. Svensk. Vetensk.-Akad. Handl., Ny Foéljd, 17 (2) : 21 (1880). The
synonym A. heteropsis Grun., the lower limit of size of the valves, and the records
from the Adriatic and Florida, and in part that from the Mediterranean, all refer
to A. fimbriata. The remainder of the account is based on A. danica.

Cleve & Mdller, Diatoms 6 : 3, n. 311 ; 4, n. 320 (1882). Both are A. fimbriata.

Cleve in Nordensk., Vega-Exped. Vetensk. Iaktt. 3 : 512 (1883). The synonym
Stauroneis cornuta Leud.-Fortm. refers to A. fimbriata. There is no information
about the identity of the specimen on which this record is based, but it is most
probably A. fimbriata, since, as explained above, Cleve later misidentified A.
fimbriata as A. danica and included A. danica in A. lorenziana.

Grunow in Van Heurck, Typ. Synops. Diat. Belg.: 121, n. 545 (1887). This is A.
Jimbriata.

Pantocsek, Beitr. Kenntn. Foss. Bacill. Ung. 1 : 31 (1886). I have seen no specimen
from Dolje to which this record might refer. Both A. fimbriata and A. baldjicki
subsp. lorenziana, but not A. danica, occur in other Hungarian fossil deposits,
and the record is probably based on one or other of these.

Pantocsek, op. cit. 2 : 57, t. 4 fig. 66 (1889). The figure is of A. baldjicki1 subsp.
lorenziana, but A. fimbriata also occurs in material from Bremia, Hungary, and
it is probable that Pantocsek did not distinguish the two species and that the
record applies to both.

De Toni, Syll. Alg. 2 : 480 (1891). The lower limits of size of the valve, and the
synonyms A. heteropsis Grun. and Stauroneis cornuta Leud.-Fortm., refer to A.
fimbriata. Of the locality records, Knarrhoi, Jutland, refers to A. danica, and so
in part do Mediterranean and Ceylon, whilst the others refer to A. fimbriata,
except that both that species and A. baldjickii subsp. lorenziana are covered by
the record from Bremia, Hungary.

A. Schmidt, Atl. Diat.: t. 198 figs. 60, 61 (1895), as A. danica var. maxima. These
figures depict A. fimbriata.

A. Schmidt, tab. cit. fig. 50 (1895). This figure of a raphe valve is probably

80 CAPARTOGRAMMA AND SCHIZOSTAURON

correctly identified as A. danica. Fig. 59 on the same plate, named by Cleve A.
lorenziana, is certainly A. danica.

Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 27 (3) : 186 (1896). The
only element in the description which refers to A. danica is the statement :
“costae [on the raphe-less valve] . . . alternating with fine lineolae, twice as close
as the costae’’. The remainder of the description, including that of A. danica
var. major Cleve, and all the synonyms except Cocconeis danica Flogel, refer to
A. fimbriata. The locality records do so also, except “‘ Mediterranean Sea ”’,
which is taken from Grunow (1880) and is based on gatherings from the Balearic
Islands in which both A. danica and A. fimbriata are represented.

H. & M. Peragallo, Diat. Mar. France : 7, t. 2 figs. 1-3 (1897). The statement
that the costae on the raphe-less valve are 8-13 in 10u (which is inconsistent
with fig. 2, correctly identified as A. danica, as also is fig. 1) and the description
of the raphe valve refer to A. fimbriata, as do the synonym A. heteropsis Grun.
and the record from “ Mer du Nord ”’, also in part that from “‘ Baléares’’. The
fig. 3 also depicts A. fimbriata.

Tempére & Peragallo, Diatomées, ed. 2 : 82, ns. 152, 153 (1908). There is no
specimen of this or any related species on either of these two slides in the set in
the British Museum Herbarium.

Tempére & Peragallo, op. cit.: 185, ns. 346-349 (1909). According to the index
(op. cit., Tables : 9 (1915)), the species is only found on ns. 348, 349. In the
set in the British Museum Herbarium there is a specimen of A. fimbriata on
n. 348 but none of this or any related species on any other of the slides. I have
found specimens of A. baldjickit subsp. lorenziana on other preparations of material
from the same fossil deposit at Bremia, near Kavna, Hungary.

Tempére & Peragallo, op. cit.: 200, n. 371 (1910). A. baldjickit subsp. lorenziana
is present on this slide in the set in the British Museum Herbarium.

Tempére & Peragallo, op. cit.: 232, ns. 442, 443 (1910). According to the index
(op. cit., Tables : g (1915)), the species is to be found on n. 443. There is
no specimen of this or any related species on this slide in the set in the British
Museum Herbarium.

Hustedt in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 396, fig. 845 (1933). The descrip-
tion is of A. danica, except that the statement that the striae in the centre of the
raphe valve are 26 in 10 would fit A. fimbriata but not A. danica ; “ 26”’ is,
however, probably an error for ‘‘ 20”’ since Hustedt’s figure of this valve shows
only 20 striae in 10 in the centre of the valve. This figure and that of the
raphe-less valve represent A. danica. The synonyms Schizostauron fimbriatum
Grun., A. heteropsis Grun., and Stauroneis cornuta Leud.-Fortm. refer to A.
fimbriata. Of the gatherings cited by Hustedt, Cleve & Moller, Diatoms, ns. 154,
155 contain A. fimbriata, Tempére & Peragallo, Diatomées, ed. 2, ns. 348, 349
contain A. fimbriata and probably also A. baldjickii subsp. lorenziana, n. 371
contains A. baldjickit subsp. lorenziana, and n. 443 probably contains one or
other or both of these species.

Cleve-Euler in K. Svensk. Vetensk.-Akad. Handl., Ser. 4, 4 (5) : 18, fig. 510 (1953).

CAPARTOGRAMMA AND SCHIZOSTAURON 81

The description of the raphe-less valve refers to A. danica but that of the raphe
valve repeats Hustedt’s error that the striae are 26 in 10 at the centre. The
only original figure, fig. 510 c, is indeterminable, but the density of its striation
and also, to some extent, its shape suggest A. fimbriata rather than A. danica.
If this is so, the fossil record from Skagganias in Smaland also refers to A. fimbriata.

Job, Suppl. 1 List Diat. Alexandria : [4] (1961). This record is based on correctly
identified specimens.

ACHNANTHES BALDJICKII (Brightw.) Grun. in K. Svensk. Vetensk.-Akad. Handl.,
Ny Foljd, 17 (2) : 21 (1880).—De Toni, Syll. Alg. 2 : 481 (1891).—Cleve in K.
Svensk. Vetensk.-Akad. Handl., Ny Foljd, 27 (3) : 187, t. 3 figs. 4, 5 (1896).

Odontidium baldjickit Brightw. in Quart. Journ. Microsc. Sci. 7 : 180, t. 9 fig. 10 (1859).

Dimeregramma baldjickii (Brightw.) Ralfs in Pritch., Hist. Infus., ed. 4 : 791 (1861).—
Walker & Chase, Not. New & Rare Diat. [1] : 1, t. 1 fig. 3 (1886).

Rhaphoneis baldjickit (Brightw.) Grun. in Verh. Zool.-bot. Ges. Wien 12 : 379 (1862).

Achnanthidium baldjickii (Brightw.) Grun. ex A. Schmidt, Atl. Diat.: t. 198 figs. 44-48
(1895).

Valves rhombic to lanceolate, sometimes slightly gibbous at the centre, subrostrate
to rostrate at the apices, 25—gou long, 13-40 broad. Raphe-less valve with a
lanceolate or moderately broad linear area, often transversely expanded on one or
both sides at the centre ; striae each consisting of a single hyaline alveolus, distant,
slightly to moderately radiate, some in the centre part of the valve usually short.
Raphe valve with an elevated marginal zone ; raphe filiform, straight except near
the central nodule, where the two halves are slightly bent in opposite directions ;
axial area narrow ; central area transversely expanded ; striae sigmoid, very
strongly radiate, longer and shorter in the central part of the valve, consisting of a

Fics. 27-29. Achnanthes baldjickii (Brightw.) Grun. subsp. baldjickii. Fig. 27, raphe
valve (in B.M. 43404) (x 1,000) ; Fig. 28, detail of same valve as Fig. 27 ( x 2,000) ;
Fig. 29, raphe-less valve of same specimen as Fig. 27 ( x 1,000).

Fics. 30-31. Achnanthes baldjickii subsp. lorenziana (Grun.) Ross. Fig. 30, raphe valve
(in Coll. Grunow n. 864a) ; Fig. 31, raphe-less valve (in Coll. F. W. Adams n. E 230).
(Both x 1,000.)

82 CAPARTOGRAMMA AND SCHIZOSTAURON

single row of fine puncta except on the elevated marginal zone, where they are
double (? and treble) rows of puncta separated by costae.

Subsp. BALDJICKU. (Figs. 27-29.)

Achnanthes baldjickti var. subquadrata Pant., Beitr. Kenntn. Foss. Bacill. Ung. 3 : t. 13
fig. 202 (1892) ; Beschr. Neu. Bacill. Pars III Beitr. Kenntn. Foss. Bacill. Ung.: 4 (1905).

Valves rhombic, usually subrostrate and gibbous at the centre, 25-goy long,
17°5-40u broad. Raphe-less valve with long and short striae in the centre part,
striae radiate, gently curved, 3-6 in 10y, closer towards the apices. Raphe valve
with shortened striae to about half-way to the apices ; striae g-12 in I0y at the
inargin, 6-7-5 in 10u against the raphe.

Known only from marine pliocene deposits in Bulgaria and Hungary.

FossiL : Phocene :

BULGARIA: Balchik (B.M. 2305, 2311, 3399, 7157, 9721, 21073, 21074, 21620 ;
B.M. 26856, 26857 = Van Heurck, Typ. Synops. Diat. Belg. ns. 545, 546; B.M.
30827, 31384, 32629, 32630, 32836, 33305, 33323, 43407, 43408, 45881, 54190, 58689 ;
Coll. F. W. Adams ns. A 167, Bess. 895, Bess. 1660, F 153 in Herb. Brit. Mus.).

HunGary : Borostelek (B.M. 33350, 70882).

Subsp. lorenziana (Grun.) Ross, stat. nov. (Figs. 30, 31.)

Rhaphonets lorenziana Grun. in Verh. Zool.-bot. Ges. Wien 12 : 381, t. 4 fig. 5 a, b (1862)
(R. fluminensis errore in expl. tab., tom. cit.: 471).—Pant., Beitr. Kenntn. Foss. Bacill.
Ung. 2 : 63, t. 28 fig. 401 (1889) (R. lorenziana var. minor in expl. tab.).

Rhaphoneis fluminensis sensu Van Heurck, Synops. Diat. Belg.: t. 36 fig. 34 (1881) ; non
Grun.

? Rhaphoneis scutellum sensu Petit in Mém. Soc. Sci. Nat. Math. Cherbourg 23 : 205, t. 3
fig. 6 (1881) ; non Ehrenb.

Achnanthes danica sensu Pant., tom. cit.: 57, t. 4 fig. 66 (1889) ; non Grun.

Achnanthes baldjickii sensu Pant., op. cit. 3 : t. 13 fig. 204 (1892) ; Beschr. Neu. Bacill.
Pars. III Beitr. Kenntn. Foss. Bacill. Ung.: 4 (1905) ; non Grun. sensu stricto.

Achnanthes lorenziana (Grun.) Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Fdéljd,
27 (3) : 186 (1896) excl. parte maxima descr.; non A. lorenziana Grun. ex A. Schmidt
(1895).—H. & M. Perag., Diat. Mar. France : 8, t. 2 fig. 4 (1897).—Hust. in Rabenh.,
Krypt.-Fl. Deutsch. 7 (2) : 395, fig. 844 (1933) excl. descr. valvae cum raphide.—Cleve-
Euler in K. Svensk. Vetensk.-Akad. Handl., Ser. 4, 4 (5): 18, fig. 511 a (1953) excl. descr.
valvae cum raphide.

Actinoneis lorenziana (Grun.) Mereschk. in Mém. Soc. Nat. Nouv.-Russ. 24 (2) : 43, 65,
figs. 12, 13 (1902) (? excl. descr., fig. et var. parva Mereschk.).

Valves rhombic-lanceolate to lanceolate, subrostrate to rostrate, 25—5ou long,
13-20 broad. Raphe-less valve with one or two short striae opposite the central
nodule on one side, usually without other short striae, sometimes with one or two
more ; striae scarcely to slightly radiate, 6-10 in I0u. + Raphe valve with shortened
striae only opposite the central area ; striae 13-16 in Iou.

Mediterranean (and possibly other seas—see below), and fossil from marine
miocene deposits in Hungary and marine pliocene deposits in Austria, Hungary
and Bulgaria.

SPAIN : Balearic Islands (Coll. F. W. Adams ns. E 230, E 231 in Herb. Brit. Mus.).

ITALy : Capri (B.M. 33068, 71125).

CAPARTOGRAMMA AND SCHIZOSTAURON 83

JuGosLaviA: Mali LoSinj, LoSinj Island, Gulf of Kvarner, Reichardt (Coll.
Grunow ns. 864a, 864b in Herb. Vienna). Porto Zubzamki, SuSak Island, Gulf of
Kvarner, Reichardt (Coll. Grunow n. 869b in Herb. Vienna).

FossiL: Miocene :

HunGary : Nyermegy (B.M. 68520, 68521 = Temp. & Perag., Diatomées, ed. 2, ns.
174, 175).

Pliocene :

AUSTRIA : Styria (B.M. 68717 = Temp. & Perag., Diatomées, ed. 2, n. 371).

Buicaria : Balchik (B.M. 2311, 7157 ; B.M. 26856, 26857 = Van Heurck, Typ.
Synops. Diat. Belg., ns. 545, 546 ; B.M. 36269, 36230, 33305).

HunGary : Bory (B.M. 33342). Bremia, near Kavna (B.M. 33302).

It is not possible to be completely certain that the striae on the raphe valve of
Achnanthes baldjickit subsp. lorenziana consist of a double row of puncta near the
margin. All the raphe valves seen have been parts of whole frustules and the
structure of the striae in the marginal zone is too fine to be resolved with the optical
microscope, at any rate in whole frustules where the presence of the other valve
interferes with image formation. Nevertheless it is possible to see that the striae
broaden as they approach the marginal zone in a way which suggests that they,
like those of the type subspecies, are double there. The costae between the striae
are certainly present in the marginal zone of both subspecies.

The very characteristic structure of the raphe valve (Figs. 27, 28, 30) of this
species has not previously been correctly described or adequately figured. The
descriptions by Grunow (K. Svensk. Vetensk.-Akad. Handl., Ny Féljd, 17 (2) : 21
(1880)) and by Cleve later in the same periodical (27 (3) : 187 (1896)) do not mention
the doubling of the puncta near the margin, nor the costae, and these two features
are not shown in Cleve’s figure (tom. cit.: t. 3 fig. 5 (1896)) nor in those of A. Schmidt
(Atl. Diat.: t. 198 figs. 45, 48 (1895)). It has already been pointed out (p. 78) that
the description of the raphe valve of A. lorenziana given by Cleve in the Synopsis
of the Naviculoid Diatoms and copied by Hustedt and Cleve-Euler is actually of
the raphe valve of A. danica.

In the pliocene deposit from Balchik, Bulgaria, in which both subspecies occur,
there are a few individuals which are to some extent intermediate between them in
that, whilst having the outline of A. baldjickii subsp. lorenziana and a raphe-less
valve with closer and less radiate striae, they have one or two shortened striae on
this valve other than the two on one side opposite the central nodule. I have
assigned such specimens to A. baldjickii subsp. lorenziana.

The two subordinate taxa within the species have very different ranges in time,
and possibly also in space, and when they do occur together in the same fossil
deposit intermediates are very few. There is thus adequate basis for distinguishing
the two as subspecies, but their close resemblance in morphology and the few
intermediates found in the Balchik material preclude separation at a higher taxonomic
level.

The type subspecies is only known from the pliocene of Balchik, Bulgaria, and of

84 CAPARTOGRAMMA AND SCHIZOSTAURON

Borostelek, Hungary. A. baldjickit subsp. lorenziana, however, occurs in the mio-
cene of Nyermegy, Hungary, and the pliocene of Bory and Bremia, both in Hungary,
of Balchik and of Styria, and it is also found in recent gatherings. I have seen
only one specimen from Bory and one broken one from Bremia, each consisting
solely of a raphe-less valve. They both have 7-7°5 striae in 10u, and there is
sufficient of the specimen from Bremia, which is broken across near the centre, to
show that at the centre it has two short striae on one side. Whilst the identifications
of these two specimens are made with confidence, there can be no absolute certainty
in the absence of associated raphe valves. A. hauckiana Grun. is present in the
Bremia material, but the specimens referred to A. baldjickit subsp. lorenziana
have their striae so widely spaced and so narrow that it is unlikely that they are
raphe-less valves of A. hauckiana.

The species is rare in recent gatherings. I have only seen it from the Gulf of
Kvarner, from Capri, and from the Balearic Islands. H. Peragallo’s (in H. & M.
Perag., Diat. Mar. France : t. 2 fig. 4 (1897)) figure of a specimen from Barcelona
is obviously correctly identified. Leuduger-Fortmorel (Bull. Soc. Bot. France 25 :
23 (1878)) records it as Rhaphoneits lorenziana from Verdelet, Cdtes-du-Nord,
France, but gives no figure or description. Cleve records A. lorenziana from Bab
el Mandeb and the Macassar Straits, but there can be no certainty as to whether
these records are correct or are based on A. danica. He also considers that Petit’s
record of Rhaphoneis scutellum from Ning-po, China, is based on this species, but
Petit’s figure is such that this identification, although possible, is doubtful. The
specimens from the Sea of Azov described and figured by Mereschkowsky as A ctinonets
lorenziana var. parva can also be referred here only with considerable doubt.
QOstrup (Danske Diat.: 126 (1910)) records A. lorenziana from Vesterhavsgerne,
Denmark, and Cleve-Euler, whose figure is a copy of Grunow’s, records it from
a marine deposit of unspecified age at Robertsfors, Sweden. There is therefore
some possibility that the species may occur recent outside the Mediterranean, but,
in view of the possibility of its being confused with others, this must remain uncertain
in the absence of specimens.

ACHNANTHES HUNGARICA (Grun.) Grun. in K. Svensk. Vetensk.-Akad. Handl., Ny
Féljd, 17 (2) : 20 (1880).—@strup, Danske Diat.: 128, t. 3 fig. 78 (1910).—Hust.
in Pascher, Siissw.-Fl. Mitteleur. 10, ed. 2 : 201, fig. 283 (1930) ; in Rabenh.,
Krypt.-Fl. Deutsch. 7 (2) : 383, fig. 829 (1933).—Cleve-Euler in K. Svensk.
Vetensk.-Akad. Handl., Ser. 4, 4 (5) : 23, fig. 521 (1953). (Figs. 32-37.)

Achnanthidium hungaricum Grun. in Verh. Zool.-bot. Ges. Wien 13 : 146, t. 4 fig. 8 (1863).

? Achnanthidium neglectum Schum. in Schr. Phys.-6kon Ges. K6nigsb. 8 : 54, t. 1 fig. 17
(1867).

Schizostauron andicola Cleve in Ofvers K. Vetensk.-Akad. Férhandl. Stockh. 38 (10) : 12,
t. 16 fig. 8 (1882) (‘‘ andicolum’’).

Stauroneis andicola (Cleve) Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Féljd, 26 (2) :
151 (1894).?

Stauroneis andicola (Cleve) M. Perag., Cat. Gén. Diat.: 829 (1897).

1 This combination, although used here by Cleve, was not validly published by him (see pp. 63-64).

CAPARTOGRAMMA AND SCHIZOSTAURON 85

Cocconets hungarica (Grun.) Schoenf., Diat. Germ,: 126, t. 13 fig. 239 (1907).

Achnanthes andicola (Cleve) Hust. in Abhandl. Naturwiss. Ver. Bremen 20 : 279, t. 2
figs. 23, 24 (1911) ; in Pascher, Siissw.-Fl. Mitteleur. 10, ed. 2 : 201, fig. 284 (1930) ;
in Rabenh., Krypt.-Fl. Deutsch. 7 (2) : 384, fig. 830 (1933).—Cleve-Euler, loc. cit.,
fig. 522 (1953) excl. B producta.

Microneis hungarica (Grun.) Meister in Beitr. Krypt.-Fl. Schweiz 4 (1) : 98, t. 13 figs. 5, 6
(1912).

Oblong-lanceolate with cuneate ends, the largest specimens sometimes gibbous
in the centre and with subrostrate apices. Raphe-less valve with very narrow, very
slightly oblique pseudoraphe and small circular central area, or, in some of the
largest specimens, with a narrow transverse fascia ; striae parallel in the centre,
radiate towards the apices, 18-20 in I0y, punctate. Raphe valve with a straight
filiform raphe with the terminal fissures turned in opposite directions ; axial area
very narrow ; central area transverse and widened outwards on one or both sides
of the valve ; large hyaline terminal areas on the largest specimens but not on
others ; striae parallel in the centre, radiate towards the apices, 20-22 in Ion,

punctate.
32
33 ‘

35
3 % 37

Fics. 32-37. Achnanthes hungarica (Grun.) Grun. (in Coll. P. T. Cleve n. 1208). Figs.
32-35, raphe valves ; Figs. 36, 37, raphe-less valves. (All x 1,000.)

Cosmopolitan in fresh waters, normally as an epiphyte on aquatic phanerogams,
especially Lemnaceae, and fossil from a freshwater miocene deposit in France.
Because of the wide distribution and frequent occurrence of this species the only
specimens cited are the type of Schizostauron andicola and the slides in published
sets in which the species occurs.

ARGENTINA : Sierra de Velasco, Prov. Rioja (Coll. P. T. Cleve n. 1208 in Herb.
Mus. Stockh., type of Schizostauron andicola).

Ecuapor : Pallatanga (B.M. 68928 = Temp. & Perag., Diatomées, ed. 2,
n. 579).

FRANCE : St. Cloud (B.M. 14625 = Temp. & Perag., Diatomées, n. 371 ; B.M.
69016 = Temp. & Perag., Diatomées, ed. 2, n. 666).

BELGIuM : Austruweel, near Antwerp (B.M. 26507 = Van Heurck, Typ. Synops.
Diat. Belg., n. 196).

86 CAPARTOGRAMMA AND SCHIZOSTAURON

Cleve records both Achnanthes hungarica and Schizostauron andicola from the
type gathering of the latter. In this gathering there are abundant specimens of
A. hungarica ranging in length from 15 to over 40ou. The tranverse fascia in the
centre of the raphe valves over 40u long and a small proportion of those slightly
shorter than this widens outwards on both sides (Figs. 32-34), but on the other
raphe valves it widens outwards on one side only (Fig. 35). The raphe valves
over 40u long have hyaline areas about 1:5 long at each apex, and most of these
valves have a rather irregular outline, with a tendency to triundulate margins
(Fig. 33). I interpret these as being post-auxospore valves of A. hungarica and
find confirmation of this view from a valve 4ou long with a hyaline area at one
apex but not the other (Fig. 34). Cleve gives the length of Schizostauron andicola
as 36u and makes no mention of hyaline areas at the apices, nor does his figure
show them. His type was accordingly one of the very few specimens with a
transverse fascia widening outwards on both sides but no hyaline areas, such as that
illustrated in Fig. 32.

In a small proportion of the raphe-less valves over 40u long there is a moderately
broad transverse fascia that widens outwards (Fig. 37) ; in the type preparation
there are appreciably fewer of these valves than of raphe valves with hyaline apices.
All the remaining raphe-less valves, even the largest, have no transverse fascia
(Fig. 36).

The material from St. Cloud distributed by Tempére and Peragallo also contains,
among abundant A. hungarica, a number of raphe valves over 40u long with hyaline
terminal areas. The corresponding raphe-less valves have a very narrow transverse
fascia, but there is no fascia on the shorter ones.

Ostrup figures three raphe valves under the name A. hungarica var. andicola ;
all have the fascia expanding outwards on both sides and the two larger have
hyaline terminal areas. The raphe-less valve figured by him under the same name
has a very narrow transverse fascia similar to that on the largest raphe-less valves
in the St. Cloud material. According to his description and figure the terminal
fissures of the raphe are both turned in the same direction, but in the Sierra de
Velasco and St. Cloud gatherings the long specimens with hyaline terminal areas,
like the shorter ones without such areas, have the terminal fissures turned in opposite
directions.

According to Hustedt’s first (1911) description and figure of A. andicola, the
terminal fissures are both bent in the same direction and the raphe-less valve has
a narrow transverse fascia. In his later accounts (1930, 1933) the direction of
the terminal fissures is neither mentioned nor shown in the figures, but the raphe-
less valve is said to have a broad fascia widening outwards like that on the raphe
valves. In a more recent paper, Hustedt (Abhandl. Naturwiss. Ver. Bremen 34 :
246 (1957)) comments on the similarity of A. andicola to A. hungarica and on the
sporadic occurrence of the former.

It thus appears that not only was Schizostauron andicola based on a large specimen
of A. hungarica but also the various diatoms later identified with Cleve’s by Ostrup
and Hustedt are post-auxospore specimens of the same species. This would account

CAPARTOGRAMMA AND SCHIZOSTAURON 87

for their sporadic occurrence, their constant association with A. hungarica, and their
comparative rarity in the gatherings in which they have been found.

Cleve-Euler maintains A. hungarica and A. andicola as separate species and com-
ments on the discrepancies in the accounts of the terminal fissures. She also suggests
that a specimen found fossil at Osterbotten might be referred to A. andicola but
belong to a distinct infraspecific taxon. Her description and figure are insufficient
for positive identification but make it plain that the specimen in question does not
belong to this species.

NAVICULA PUPULA Kiitz., Kies. Bacill.: 93, t. 30 fig. 40 (1844).—Grun. in K. Svensk.
Vetensk.-Akad. Handl., Ny Foljd, 17 (2) : 45, t. 2 fig. 53 (1880).—Cleve in K.
Svensk. Vetensk.-Akad. Handl., Ny Foljd, 26 (2) : 131 (1894).—Hust. in Pasch.,
Siissw.-F]. Mitteleur. 10, ed. 2 : 281, fig. 467 (1930) ; in A. Schmidt, Atl. Diat.:
t. 396 figs. 10-33 (1934) ; in Internation. Rev. Hydrobiol. Hydrogr. 42 : 64
(1942) ; in Rabenh., Krypt.-Fl. Deutsch. 7 (3) : 120, figs. 1254, 1255 (1961).—
Cleve-Euler in K. Svensk. Vetensk.-Akad. Handl., Ser. 4, 4 (5) : 186, fig. 890 (1953).

Stauronets rectangularis Greg. in Quart. Journ. Microsc. Sci. 2 : 99, t. 4 fig. 17 (1854).

Navicula laevissima sensu W. Smith, Synops. Brit. Diat. 2 : 91 (1856) ; non Kiitz.—Grun.
in Verh. Zool.-bot. Ges. Wien Io : 549, t. 4 fig. 5 (1860).—Rabenh., Fl. Eur. Alg. 1 : 188
(1864).—O’Meara in Proc. R. Irish Acad., Ser. 2, 2, Sci.: 376, t. 31 fig. 54 (1875).—Brun,
Diat. Alp. Jura : 68, t. 7 fig. 32 (1880).

Navicula granum Schum. in Schr. Phys.-dkon. Ges. K6nigsb. 8 : 59, t. 2 fig. 46 (1867).

Stauroneis verbania De Not. in Erb. Critt. Ital., Ser. 2 : n. 434 (1871).

Stauroneis tatrica Gutw. in Sprawozd. Kom. Fizyjogr. 25 : (24), t. 1 fig. 20 (1890).

Schizostauron tatricum (Gutw.) De Toni in Nuova Notarisia [1] : 196 (1890).

Schizostauron verbanium (De Not.) De Toni, Syll. Alg. 2 : 225 (1891).

Sellaphora pupula (Kiitz.) Mereschk. in Ann. & Mag. Nat. Hist., Ser. 7, 9 : 187, t. 4 figs. 1-5
(1902).

Navicula pseudopupula Krasske in Bot. Arch. 3 : 197, fig. 4 (1923).—Cleve-Euler, tom.
cit.: 187, fig. 891 (1953).

Navicula pseudopupula var. aqueductae Krasske, loc. cit., fig. 8 (1923).

Navicula aqueductae (Krasske) Krasske in Abhandl. & Ber. Ver. Naturk. Cassel 54 : 44,
t. 2 fig. 23 (1925).

Navicula mutata Krasske in Bot. Arch. 27 : 354, fig. 16 (1929).

For description and figure, see Hustedt (1930, 1961).

A cosmopolitan freshwater species, and widespread in pleistocene fossil deposits.

GERMANY : Nordhausen, Kiitzing (B.M. 17918 = Coll. Kiitzing n. 58, ? type).
River Main, Miindung des [illegible] bach in dem Main, 11 Sept. 1847, A. Braun
(B.M. 18726 = Coll. Kiitzing n. 925).

FRANCE : Falaise, Brébisson (B.M. 18725 = Coll. Kiitzing n. 1468).

ENGLAND : Lewes, Sussex, Mar. 1853, W. Smith (B.M. 23638).

ScoTLAND : Lochend, near Edinburgh, 7 Mar. 1854, Gregory (B.M. 20507, 48023).

ITaLy : Verbania, in a puddle near the mouth of the San Bernardino, autumn
1868, De Notaris (Erb. Critt. Ital., Ser. 2, n. 434, isotype of Stauronets verbanta).
FossIL : Pleistocene :

ScoTLAND : Knock, Isle of Mull, Duke of Argyll (B.M. 22320, isotype of Stauronets
rectangularis).

88 CAPARTOGRAMMA AND SCHIZOSTAURON

The outline of this species is very variable and there is quite a wide range in
the density of its striation. In consequence it is usually regarded as having many
varieties (cf. Hustedt, 1930, 1961), or as consisting of more than one species (cf.
Cleve-Euler, 1953). When a large number of populations are examined, however,
the various forms are found so to intergrade that there seems little justification for
the taxonomic recognition of them at any level.

In his protologue Kiitzing gives the locality of this species as “‘ Lebend in siissem
Wasser bei Nordhausen’’. The only specimens in his herbarium labelled Navicula
pupula are the one from the River Main collected by A.Braun and the one from
Falaise collected by Brébisson which are cited above. Both these came into his
possession after the publication of the Bacillarien. Accordingly, he must either
have failed to preserve the gathering in which he found the species or have found
it in one of the gatherings from Nordhausen which are preserved in his herbarium
but have failed to record the identification on the label. I have examined these
gatherings and on his n. 58 I found small numbers of NV. fupula as currently under-
stood. These match his figure well, better than anything else present in any of
his gatherings from Nordhausen. This gathering is labelled ‘‘ Fragilaria corrugata
Kg. Nordhausen ”’ in Kiitzing’s hand and is the only one of that species in his
herbarium labelled with its name. In his protologue of Fragilaria corrugata Kiitzing
(Kies. Bacill.: 45 (1844)) says “In Siisswassergraben unter verschiedenen andern
Diatomeen bei Nordhausen ’’.

Further confirmation that the current concept of N. pupula corresponds to
Kiitzing’s is provided by the two gatherings in his herbarium labelled with its
name. Both these contain the species as now understood. The identification on
the A. Braun specimen is in Kiitzing’s own hand. That on the Brébisson specimen
is written in ink by Brébisson, but other identifications have been added in pencil
in Kiitzing’s hand, and the words “ Navicula Pupula Kg.”’ have been underlined
in pencil and the full stop after ‘“ Kg’ converted into an exclamation mark by a
pencil stroke, presumably by Kiitzing.

This species was independently described by Gregory in 1854 as Stauroneis
rectangularis, and W. Smith in 1856 applied the name Navicula laevissima Kiitz.
to it, citing S. rvectangularis asa synonym. W.Smith’s Lewes gathering and Gregory’s
from Lochend, together with the original material of S. rectangularis, represent
such of the material on which W. Smith’s account of N. laevissima is based as is
preserved in the British Museum Herbarium. Until 1880 most authors followed
W. Smith and applied the name N. laevissima to this species. In 1860, however,
Grunow suggested that N. pupula might be this species, and in 1880, after seeing
“specimina authentica ’’, he stated firmly that this was so. At the same time he
expressed doubts about the true identity of N. laevissima, and ever since then it

1 At this time Grunow was collaborating with Van Heurck in the preparation of the latter’s Synopsis
des Diatomées, and Van Heurck had acquired at Eulenstein’s death a duplicate set of Kiitzing’s diatoms
removed from the original packets by Eulenstein when the collection was in his hands (cf. Miller &
Zaunick, Friedrich Traugott Kiitzing 1807-1893 Aufzeichnungen & Erinnerungen : 20 (1960)). The
authentic material Grunow saw was therefore probably some of A. Braun’s River Main gathering

and certainly Brébisson’s Falaise gathering, for his figure of the species is of a specimen from “ Falaise,
von Kiitzing selbst bestimmt ”’.

CAPARTOGRAMMA AND SCHIZOSTAURON 89

has been treated as a doubtful species (cf. Hustedt in Pascher, Siissw.-Fl. Mitteleur.
10, ed. 2 : 279 (1930)). Its type, however, shows it to be the species normally
known as N. bacilliformis Grun., for which it provides the correct name (see p. 90
below).

This species is the type of the genus Sellaphora Mereschk. (Ann. & Mag. Nat.
Hist., Ser. 7, 9 : 186 (1902)). There is little doubt that it and the type of Navicula
Bory belong to different genera, but until a comprehensive review of this group of
genera can be undertaken, it seems best to retain it in its currently accepted position.

One of the names included by Hustedt (1961) in the synonymy of N. pupula is
N. borscowit Mereschk. That combination has never been validly published ; at
the place referred to, Mereschkowsky described Sellaphora borscow1i as a new species
based on marine specimens from San Pedro, California, that lived in Schizonema-
type tubes. There is nothing in his description or figure (Ann. & Mag. Nat. Hist.,
Ser. 7, 9 : 188, t. 4 figs. 6-10 (1902)) to suggest that it is this species.

As Mayer (Denkschr. Regensb. Bot. Ges. 21 : 168 (1940)) points out, Krasske’s
original descriptions and figures of his N. pseudopupula, N. aqueductae and N.
mutata do not show the characteristic terminal nodules of this species, nor, in the
case of N. mutata, the laterally expanded central area. Hustedt’s figures in A.
Schmidt’s Adlas (t. 396, figs. 10-14, 29-31 (1934)) of Krasske’s original material,
however, show these features. Krasske attributes the figures of N. pseudopupula
and N. pseudopupula var. aqueductae accompanying his original accounts of these
taxa to Hustedt, so there is every reason for believing that Hustedt’s later figures
are of the same taxa. He has explained (in Pascher, Siissw.-Fl. Mitteleur. 10, ed.
2 : 282 (1930) ; Internation. Rev. Hydrobiol. Hydrogr. 42 : 65 (1942)) that the
original failure to see the characteristic features of these diatoms was due to the use
of a mounting medium of too low a refractive index.

NAVICULA LAEVISSIMA Kiitz., Kies. Bacill.: 96, t. 21 fig. 14 (1844).
Stauroneis wittrockit Lagerst. in Bih. K. Svensk. Vetensk.-Akad. Handl. 1 (14) : 38, t. 2
fig. 15 (1873).
Navicula bacilliformis Grun. in K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 17 (2) : 44,
t. 2 fig. 51 (1880).—Hust. in Pasch., Siissw.-Fl. Mitteleur. 10, ed. 2 : 273, fig. 446 (1930) ;
in A. Schmidt, Atl. Diat.: t. 396, figs. 7-9 (1934).—Cleve-Euler in K. Svensk. Vetensk.-
Akad. Handl., Ser. 4, 4 (5) : 188, fig. 892 (1953).
Navicula fusticulus Ostr., Danske Diat.: 36, t. 1 fig. 19 (1910).
Navicula wittrockii (Lagerst.) Cleve-Euler in Comment. Biol. Soc. Sci. Fenn. 4 (14) : 86,
t. 5 fig. 145 (1934) ; in K. Svensk. Vetensk.-Akad. Handl., Ser. 4, 4 (5) : 188, fig. 893
(1953).—Hust. in Rabenh., Krypt.-Fl. Deutsch. 7 (3) : 124, figs. 1256, 1257 (1961).
For description and figure, see Hustedt (1930 as N. bacilliformis, 1961 as N.
wittrockit).
A cosmopolitan freshwater species, and widespread in pleistocene fossil deposits.
FossiIL: Pleistocene :
ITALY : Santa Fiore (B.M. 18747, 18748 = Coll. Kiitzing n. gb ; B.M. 18820 =
Coll. Kiitzing n. gII ; isotypes).

This species is based on specimens that were found “ fossil im Bergmehl von San
Fiore’. Among the specimens in the British Museum Herbarium labelled “ Coll.

fol) CAPARTOGRAMMA AND SCHIZOSTAURON

Kitz. Diat.”” there are a number of packets labelled in Eulenstein’s hand with a
species name, the locality ‘‘ St. Fiore’”’ and the number “ g1r’’. One of these is
labelled “‘ Navicula laevissima Kg.” and the slide n. B.M. 18820 was prepared from
this material. In other cases where such packets occur, Kiitzing’s original packet
with the same number is present under another species and bears the name of more
than one species. No original packet of n. gII is present and this number is too high
for a packet that was in Kiitzing’s possession at the time when he wrote the Baczl-
larien. There is, however, a specimen labelled “ St. Fiora Campylodiscus Eunotia
granulata Nav. binodis gb”’ in Kiitzing’s hand, and this packet contains considerable
quantities of the diatomite. It is clearly part of Kiitzing’s original sample of this
fossil deposit. Comparison of the microscope slides made from the material in
this packet with those made from the packets numbered g11 by Eulenstein shows
that all were prepared from the same fossil sample. Whether Kiitzing received a
second batch of it at some stage or whether Eulenstein mis-read Kiitzing’s ‘‘ gb ”’
as “‘ gti ”’ (this is possible since Kiitzing’s figures are written very small) one cannot
now tell. In this material the species under discussion here, i.e. that called Navicula
bacilliformis by all authors from 1880 until 1961 and N. wittrocki by Hustedt in
1961, is not uncommon and is the only one matching Kiitzing’s description and
figure. N. pupula also occurs, but much more rarely, and the specimens of it have
not the gibbous centre mentioned in Kiitzing’s description (‘‘ medio leviter
ventricosa ’’) and shown in his figure.

The name UN. laevissima appears never to have been correctly applied by any
subsequent author. W. Smith (Synops. Brit. Diat. 2 : 91 (1856)) used it for N.
pupula and in this he was followed by a number of authors (cf. p. 88 above). Donkin
(Nat. Hist. Brit. Diat.: 28, t. 5 fig. 2 (1871)), however, used it for a form which
probably belongs to N. bacillum Ehrenb. In 1880 Grunow (K. Svensk. Vetensk.-
Akad. Handl., Ny Foljd, 17 (2) : 45 (1880)), who had seen authentic material of
N. pupula but not of N. laevissima, gave an accurate account of N. pupula but
considered the identity of N. Jaevissima doubtful ; he gave a description under the
name but with a query, and further down the same page said that N. laevissima
might be identical with N. ventricosa sensu Donkin (= Calonets stlicula var. ventricosa
Cleve), and that the species he had described under the name N. Jaevissima should
be given the new name N. pseudobacillum. At the same time he described N.
bacilliformis as a new species and suggested that N. laevissima sensu Donkin might
belong to it. In Van Heurck’s Synopsis, published later in the same year, Grunow
was responsible for two figures on the same plate, one labelled N. pseudobacillum
(t. 13 fig. 9) and the other N. laevissima (t. 13 fig. 13). According to Hustedt (in
Pasch., Siissw.-Fl. Mitteleur. 10, ed. 2 : 280 (1930)), N. pseudobacillum is a synonym
of N. bacillum Ehrenb., and both these figures in the Synopsis seem to represent that
species.

Except for this uniting of N. bacillum and N. pseudobacillum, Grunow’s treatment
of these species in the Avrctischen Diatomeen has normally been followed until very
recently ; the name N. pupula has been correctly applied, the identity of N.
laevissima has been regarded as doubtful, and the species under discussion has been

CAPARTOGRAMMA AND SCHIZOSTAURON 91

called N. bacilliformis. H. and M. Peragallo (Diat. Mar. France : 67, t. 8 fig. 25
(1897)), however, used the name N. Jaevissima for a marine form quite unrelated to
any of the species to which it had previously been applied, or misapplied, and
Pantocsek (Res. Wiss. Erforsch. Balatonsees 2 (2, 1, Anhang) : 47, t. 5 fig. 112 (1902))
used the name in yet another sense.

In 1961 Hustedt called this species N. wittrockii (Lagerst.) Cleve-Euler since he
considered that the type material of Stawroneis wittrockii Lagerst. was conspecific
with the type material of N. bacilliformis, an opinion which seems to me to be justified
by the original accounts of the two. Cleve-Euler herself, on the other hand,
maintains that there are two species involved and that N. wittrockii is distinguished
from N. bacilliformis by having straight striae.

Navicula caucasica Ross, nom. nov. (Fig. 38.)
Schizostauron rhombicum M. Perag. in Temp. & Perag., Diatomées, ed. 2 : 398, n. 825 (1913).

Valves narrowly rhombic-lanceolate with obtuse, rounded apices, 47-5—60y long,
12-15u broad. Raphe slightly sinuous, somewhat oblique, central pores close,
terminal fissures hooked, both turned in the same direction. Axial area rather
narrow. Central area three-quarters of the breadth of the valve, expanded out-
wards. Terminal areas prominent and rather large. Striae radiate at the centre,
convergent at the apices, somewhat curved, very finely punctate, 14-15 in Ioy ;
short striae opposite the central nodule of uneven length.

38
Fic. 38. Navicula caucasica Ross (in B.M. 69176) (x 1,000).
Known from a single freshwater gathering from the Caucasus.
U.S.S.R.: Kisstib, Caucasus (B.M. 69176 = Temp. & Perag., Diatomées, ed.
2, n. 825, isotype).
This species resembles Stauroneis amphioxys Greg. but the central area does not

extend so far across the cell and widens outwards more markedly, and the striae
are wider spaced, more strongly radiate at the centre, and convergent at the apices.

92 CAPARTOGRAMMA AND SCHIZOSTAURON

In spite of this resemblance, I place this species in Navicula, not Stauronets, since
the central area is not a stauros and the striae do not appear under the light
microscope to have the structure typical of true Stawroneis.

STAURONEIS SMITHII var. SAGITTA (Cleve) Hust. in Rabenh., Krypt.-Fl. Deutsch. 7
(2) : 811, fig. 1158 (1959).

Pleurostauron sagitta Cleve in Cleve & Mller, Diatoms 5 : 6, n. 261 (1879), nom. nud.

Stauroneis sagitta Cleve in K. Svensk. Vetensk.-Akad. Handl., Ny Foljd, 18 (5) : 15, t. 3
fig. 45 (1881) ; op. cit. 26 (2) : 151 (1894).—Cleve-Euler in K. Svensk. Vetensk.-Akad.
Handl., Ser. 4, 4 (5) : 217, fig. 960 (1953).

Stauroneis stefanssonii Ostr. in Rosenv. & Warm., Bot. Iceland 2 : 15, t. 2 fig. 18 (1918)
(‘‘ Stefanssoni’’).

For description and figure, see Hustedt (loc. cit.).

From fresh and slightly brackish waters in northern Scandinavia and Iceland,
and fossil from freshwater quaternary deposits in Sweden and Finland.

Norway : Mouth of the Tana Elf (B.M. 13024 = Cleve & Moller, Diatoms n. 261,
isotype).

In the Synopsis of the Naviculoid Diatoms Cleve includes Stauroneis sagitta in the
subdivision Schizostauron of the Naviculae microstigmaticae. The resemblance
between the species to which this diatom is here referred and Capartogramma is
discussed earlier in this paper (p. 53). It has been regarded as a distinct species by
most authors, but Hustedt’s treatment of it as a variety of S. smithii seems justified.
The division of the stauros on one or both sides in many specimens is the reason
why Cleve referred it to Schizostauron, but this division of the stauros is comparable
to that of a single tigillum sometimes found in Capartogramma and not to the
possession of two diverging tigilla, which is the characteristic of that genus.

Bull. B.M. (N.H.) Bot. 3, 2 PLATE 1

C

Stereo-pairs of electron micrographs: A, Capartogramma karstenii
(Zanon) Ross (x 2,500); B, C. crucicula (Grun., ex Cleve) Ross
(x 5,500); C, C. amphoroides Ross (x 2,500). Phot. K. Little.

Bull. B.M. (N.H.) Bot. 3, 2 PLATE 2

B C

A. Capartogramma crucicula (Grun. ex Cleve) Ross; stereo-pair of
electron micrographs. (x 13,000.) B,C. C. karstenii (Zanon) Ross ;
electron micrographs. (B x 50,000; C x 35,000.) Phot. K. Little.

ANGIOSPERMS OF THE
CAMBRIDGE ANNOBON
ISLAND EXPEDITION

A. W. EXELL

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

- BOTANY , Vol. 3 No. 3
| LONDON : 1963

ANGIOSPERMS OF THE CAMBRIDGE
ANNOBON ISLAND EXPEDITION

}

BY

A. W. EXELL

Pp. 93-118 ; Plates 3-12

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 3 No. 3
LONDON : 1963

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), ¢stituted in 1949, 1s
issued in five series corresponding to the Departments
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This paper is Vol. 3, No. 3 of the Botany series.

© Trustees of the British Museum, 1963

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Issued September, 1963 _ Price Fifteen Shillings

ANGIOSPERMS OF THE CAMBRIDGE
ANNOBON ISLAND EXPEDITION
By A. W. EXELL

THE following is an account of the more interesting Angiospermae collected on the
Cambridge Annobon Island Expedition 1959. The two collectors were Fenella
A. Melville (now Mrs. Wrigley) and T. C. Wrigley.

I have included in the list all new records for Annobon and specimens of species
already recorded for the island but meriting inclusion for one reason or another.
It is particularly interesting to be able to record a number of gatherings of species
described from Mildbraed’s 1g11 collection of which the types (formerly in the
Berlin Herbarium) have been destroyed. These have largely been identified ex
descriptione : if correctly named they provide (allowing for the very small size of
the island) topotypes of the species concerned. Photographs of three of the
Mildbraed types and seven of the new topotypes are included in this account as
illustrations. Except for the Orchidaceae (which are at Kew) all the specimens
collected on the Cambridge Expedition are represented in the British Museum
Herbarium, and all specimens cited in this account are in the British Museum
Herbarium unless otherwise indicated.

In my statements of new records “the islands’’ means the three islands of
Principe, S. Tomé and Annobon in the Gulf of Guinea. The four principal works
which I have already published on the flora of these islands are: (1) Catalogue of
the Vascular Plants of S. Tomé (with Principe and Annobon) (1944) ; (2) Supplement
to the Catalogue of the Vascular Plants of S. Tomé (with Principe and Annobon) (1956) ;
(3) Aditamento a Flora das Ithas de Séio Tomé e do Principe (Confer. Internacion. Afr.
Oc., Sess. 6, 3 : 77-91) (1958), with A. Rozeira ; (4) Additions to the Flora of S. Tomé
and Principe (Bull. Inst. Frang. Afr. Noire 21, Sér. A, n. 2) (1959). They are
abbreviated in this account as follows : (1) Exell, Cat.; (2) Exell, Suppl.; (3) Exell,
Aditam.; (4) Exell, Addit. The third of these works was published without my
seeing a proof and is marred by typographical errors. I have taken the opportunity
to bring the nomenclature up to date as regards a few species listed in these works
and to correct various errors which have been pointed out to me, but I have not
troubled to correct the fairly obvious orthographical mistakes in the Aditamento.
Taxa not recorded for Annobon are placed within square brackets.

I am indebted as usual to numerous colleagues at the British Museum (Natural
History), the Royal Botanic Gardens, Kew, and other institutions ; and more
particularly to J. E. Dandy (general arrangement), Miss D. Hillcoat (Leguminosae),
F. N. Hepper (Rubiaceae), J. Lewis (Loganiaceae), W. T. Stearn (Solanaceae,
Piperaceae, Loranthaceae), J. Léonard (Euphorbiaceae), V. S. Summerhayes
(Orchidaceae) and W. D. Clayton (Gramineae).

96 ANGIOSPERMS OF ANNOBON ISLAND

DILLENIACEAE
TETRACERA ALNIFOLIA Willd.—Exell, Cat.: 100.
ANNOBON: S.E. of Santa Cruz, 450 m., forest on ridge, scabrid climber and
scrambler, 7 Aug. 1959, Wrigley 249.
New record for Annobon.

ANNONACEAE
ANNONA MURICATA L.—Exell, Cat.: ror.
ANNOBON : Near Ambo, 27 m., open woodland, 20 July 1959, Wrigley 63.
New record for Annobon. Introduced. Soursop.

ANNONA RETICULATA L.—Exell, Cat.: ror.
ANNOBON : Near Ambo, 27 m., and near Crater Lake, open woodland, 16 July

1959, Wrigley 42.
New record for Annobon. Introduced. Custard-apple.

[CRUCIFERAE]
[RorIPPA INDICA (L.) Hiern, Cat. Afr. Pl. Welw. 1: xxvi (1896).—Hara in Journ.
Jap. Bot. 30 : 197 (1955).
Sisymbrium indicum L., Syst. Nat., ed. 12, 2: 441 et Mant. Pl.: 93 (1767).
Nasturtium sinapis (Burm. f.) O. E. Schulz.—Exell, Suppl.: 10 ; Aditam.: 81.
Rorippa sinapis (Burm. f.) Ohwi & Hara.—Exell, Addit.: 450.

S. TomE : Ribeira Peixe, Rozetva 236 (Herb. Porto Univ.). Diogo Vaz, Rozeira
869 (Herb. Porto Univ.).

This introduced species has previously been recorded from S. Tomé under the
names Nasturtium sinapis and Rorippa sinapis. It is widely distributed in south-
east Asia from Japan and China to India and Malaysia.

The name R. indica has been misapplied to another species, R. madagascariensis
(DC.) Hara (tom. cit.: 198).]

CAPPARACEAE
CLEOME RUTIDOSPERMA DC., Prodr. I: 241 (1824).
Cleome ciliata Schumach.—Exell, Cat.: 105 ; Suppl.: Io.

ANNOBON: N.W. of the Island, 6 m., rocks and cassava plantations by sea,
11 July 1959, Melville 129.

De Candolle’s name replaces that of Schumacher. The species is recorded for
Principe, S. Tomé and Annobon and is widespread in tropical Africa southwards
as far as Northern Rhodesia.

PORTULACACEAE
PORTULACA OLERACEA L.—Exell, Cat.: 110 ; Addit.: 450.
ANNOBON : Ambo, by the expedition’s houses, 15 m., sandy place by house,
21 July 1959, Melville 168.
New record for Annobon.

ANGIOSPERMS OF ANNOBON ISLAND 97

TALINUM TRIANGULARE (Jacq.) Willd.—Exell, Cat.: 110 ; Suppl.: 11.
ANNOBON:: N.W. of San Pedro, forest, 25 July 1959, Melville 194.
New record for Annobon. Introduced.

MALVACEAE
SIDA RHOMBIFOLIA L.—Exell, Cat.: 115.
ANNOBON : Near Capelle San Juan, 30 m., wayside in cultivated land, 10 Aug.
1959, Melville 258.
New record for Annobon.

GOSSYPIUM BARBADENSE L.—Exell, Cat.: 119.
ANNOBON:: Near Ambo, 9 m., pathside, 11 July 1959, Wrigley 12.
New record for Annobon. Introduced ; presumably an escape from cultivation.

RUTACEAE
CITRUS AURANTIFOLIA (Christm.) Swingle.—Exell, Cat.: 131.
ANNOBON : Near Ambo, 24 m., waste ground, 12 July 1959, Wrigley 25.
New record for Annobon. Introduced. Lime.

OCHNACEAE
OcHNA MEMBRANACEA Oliv., Fl. Trop. Afr. 1: 316 (1868).—Henriq. in Bol. Soc.
Brot. 10 : 105 (1893).
Diporochna quintasii Tiegh. in Ann. Sci. Nat., Sér. 8, Bot. 16 : 392 (1902).
Ochna quintasii (Tiegh.) Exell, Cat.: 132.
Ochna cf. gilgiana, Exell, Cat.: 132.
ANNOBON : Between Capelle San Juan and San Pedro, 45 m., steep wood, cliffs
by sea, 11 Aug. 1959, Melville 260.
Recorded from Annobon by Mildbraed as O. cf. gilgiana. Also in S. Tomé and
widespread in western tropical Africa, eastwards to Uganda and southwards to
Angola.

MELIACEAE
TURRAEA GLOMERULIFLORA Harms.—Exell. Cat.: 136. (Plate 3.)
ANNOBON : Ridge west of Crater Lake, 360 m., open woodland ; ‘ small sepals ;

green petals ; white filament tube with 10 stamens’; 18 July 1959, Wrigley 52.

Endemic to Annobon. Wrigley’s locality is quite near to that of the holotype,
Mildbraed 6487 (Herb. Berlin, destroyed). A photograph of the holotype exists
in the British Museum Herbarium and is reproduced in Plate 3.

CELASTRACEAE
Maytenus annobonensis (Loes. & Mildbr.) Exell, comb. nov. (Plate 4.)

Gymnosporia annobonensis Loes. & Mildbr. in Notizbl. Bot. Gart. & Mus. Berl.-Dahl. 12:
30 (1934).—Exell, Cat.: 138.

98 ANGIOSPERMS OF ANNOBON ISLAND

ANNOBON : Ridge west of Crater Lake, 360 m., open woodland ; “ flower white ;
3-merous fruit with I orange seed’; 18 July 1959, Wrigley 56.

Endemic to Annobon. This material was collected in almost the same locality
as the holotype, Mildbraed 6541 (Herb. Berlin, destroyed), and the only doubt
about the identification lies in the fact that the original specimen had neither flowers
nor fruit so that here “ex descriptione’’ has little meaning. Actually this new
gathering by Wrigley confirms the supposition that the species belongs to the genus
Maytenus (Gymnosporia). It differs from M. monodii Exell (endemic to S. Tomé)
by its dense ferrugineous pubescence on the young stems, being nearer in this
respect to M. welwitschiana Exell & Mendonca (Angola) but with larger flowers
than in the latter species and with a glabrous (or only minutely pubescent) capsule.

ANACARDIACEAE
MANGIFERA INDICA L.—Exell, Cat.: 145.
ANNOBON : Common tree, lower grassland and up to Crater Lake, regenerating
very freely, 19 July 1959, Wrigley 6r.
New record for Annobon. Introduced.

CONNARACEAE
AGELAEA OVALIS Schellenb.— Exell, Cat.: 148. (Plate 5.)
ANNOBON : Ambo, go m.; dry valley ; “tree c. 30 ft.; fruit red and yellow

with black and orange seed ; flowers off-white (pinkish) ’’; 2 Aug. 1959, Melville 232.

Endemic to Annobon. Melville’s plant is almost certainly A. ovalis, of which
the holotype, Mzldbraed 6679 (Herb. Berlin), has been destroyed. It is, however,
difficult to believe that the plant is a tree and I suspect an error in the collector’s
note.

LEGUMINOSAE
ARACHIS HYPOGAEA L.—Exell, Cat.: 155.
ANNOBON : Cultivated on a small scale during the wet season, 18 Aug. 1959,
Wrigley 297, fruit only.
New record for Annobon. Introduced.

DESMODIUM RAMOSISSIMUM G. Don.—Exell, Suppl.: 16.
Desmodium mauritianum sensu Exell, Cat.: 158.
ANNOBON : Path towards Pico do Fogo and Crater Lake, 90 m., pathside,
10 July 1959, Melville ro5.
New record for Annobon.

MUCUNA SLOANEI Fawc. & Rendle in Journ. of Bot. 55 : 36 (1917).—Hepper in
Hutch.:& Dalziel, Fl. W: Trop. Afr.,-ed..2, 1: 561 (1958):
Mucuna urens sensu Exell, Cat.: 160.
ANNOBON : Near Ambo, 24 m., climbing over trees, 14 July 1959, Wrigley 3o.
Recorded for S. Tomé and Annobon as M. urens ; widespread in tropical Africa
and tropical America. Calabar Bean.

ANGIOSPERMS OF ANNOBON ISLAND 99

VIGNA UNGUICULATA (L.) Walp., Repert. Bot. Syst. 1: 779 (1842).—Hepper in
Hutch. & Dalziel, Fl. W. Trop. Afr., ed. 2, 1 : 569 (1958).

Dolichos unguiculatus L., Sp. Pl. 2: 725 (1753).
Clitoria alba G. Don, Gen. Syst. 2 : 215 (1832).
Vigna alba (G. Don) Planch. ex Baker f.—Exell, Cat.: 161.
ANNOBON: East path towards Crater Lake, 135 m., pathside and cassava
plantation, running over rocks, 18 July 1959, Melville 162.
New record for Annobon. The species occurs also in S. Tomé and is widespread
in tropical and subtropical regions.

CAJANUS CAJAN (L.) Millsp.—Exell, Cat.: 163.
ANNOBON: Near Ambo, 30 m., open grassland, 23 July 1959, Wrigley 76.
New record for Annobon. Introduced.

ERIOSEMA GLOMERATUM (Guillem. & Perrott.) Hook. f. in Hook., Niger Fl.: 313

(1849).
Rhynchosia glomerata Guillem. & Perrott. in Guillem., Perrott. & Rich., Tent. FI.
Senegamb. I : 216 (1832).
ANNOBON : Near Ambo, 30 m., pathside, 17 July 1959, Wrigley 46.
New record of the genus Eviosema (DC.) Desv. for the islands. EF. glomeratum
is widespread in tropical Africa.

CAESALPINIA PULCHERRIMA (L.) Swartz.—Exell, Cat.: 165.
ANNOBON : Mission, Ambo, 15 m., garden, 4 Aug. 1959, Melville 233.
New record for Annobon. Cultivated. Barbados Pride.

CASSIA OCCIDENTALIS L.—Exell, Cat.: 166.

ANNOBON : Path between Capelle San Juan and San Pedro, 60 m., pathside,
2 Aug. 1959, Wrigley 230.

New record for Annobon.

CASSIA OBTUSIFOLIA L., Sp. Pl. : 377 (1753).—Brenan in Kew Bull. 13 : 248 (1958).
Cassia tora sensu Exell, Cat.: 167.

ANNOBON : Path between Capelle San Juan and San Pedro, 60 m., pathside,
2 Aug. 1959, Wrigley 230a.

New record for Annobon. The species occurs also in S. Tomé and is widespread
in the tropics.

According to Brenan (tom. cit.: 248-252) C. tora L. is a different species which
does not occur in West Africa.

CRASSULACEAE
KALANCHOE CRENATA (Andr.) Haw.—Exell, Cat.: 173 ; Suppl.: Ig.
ANNOBON : Near shore near Ambo, 6 m., edge of dry river, 4 Aug. 1959, Wrigley
235. |
New record for Annobon.

100 ANGIOSPERMS OF ANNOBON ISLAND

RHIZOPHORACEAE

CASSIPOUREA ANNOBONENSIS Mildbr. ex Alston.—Exell, Cat.: 174. (Plate 6.)

ANNOBON : Pico do Fogo on S.W. side, 360 m., edge of forest at base of Pico ;
“flowers 4-merous ; tree 30 ft. (branches well spaced for climbing)”; 25 July
1959, Melville rgo.

Endemic to Annobon. The locality is near to that of the holotype, Mildbraed 6511
(Herb. Berlin, destroyed). A photograph of the holotype exists in the British
Museum Herbarium and is reproduced in Plate 6.

COMBRETACEAE
TERMINALIA CATAPPA L.—Exell, Cat.: 175.
ANNOBON: Near Ambo, 24 m., planted; ‘the nuts are eaten’; 18 Aug.

1959, Wrigley 296.
New record for Annobon. Introduced. Indian Almond.

MELASTOMATACEAE
TRISTEMMA OREOTHAMNOS Mildbr.—Exell, Cat.: 178. (Plate 7.)
ANNOBON : Pico Surcado, 420 m., forest ; “‘ flower pale pink’; 21 July 1959,

Wrigley 65. Crater Lake, N.E. side, 360 m., forest near base of Pico do Fogo ;
“corolla pink ; anthers yellow’; 25 July 1959, Melville 193.

Endemic to Annobon. The holotype of this species, Mildbraed 6677 (Herb.
Berlin), has been destroyed ; it was collected at a somewhat higher altitude on the
neighbouring peak of Quioveo.

PUNICACEAE
PUNICA GRANATUM L., Sp. Pl. 1: 472 (1753).
ANNOBON : Ambo, near the Mission, 15 m., cultivated, 2 Aug. 1959, Melville 219.
New record of the genus Pumica L. for Annobon. P. granatum, the Pomegranate,
is a widely cultivated species of Asiatic origin.

CARICACEAE
CARICA PAPAYA L.—Exell, Cat.: 183.
ANNOBON : Crater Lake, 265 m., lakeside cultivation, 24 July 1959, Melville 182.
New record for Annobon. Introduced.

CUCURBITACEAE
[PEPONIUM VOGELII (Hook. f.) Engler in Engler & Prantl, Nat. Pflanzenfam.,
Nachtr. zum 2-4 : 318 (1897).—Jeffrey in Kew Bull. 15 : 359 (1962).
Peponia vogelii Hook. f. in Oliv., Fl. Trop. Afr. 2 : 526 (1871).
Peponium bracteatum (Cogn.) Cogn.—Exell, Cat.: 184.
Described from S. Tomé as Peponza bracteata Cogn., but this is now referred by
Jeffrey (loc. cit.) to Pepbonium vogelii, a species widely distributed in tropical Africa
and extending into Natal.]

ANGIOSPERMS OF ANNOBON ISLAND IOI

LAGENARIA BREVIFLORA (Benth.) Roberty in Bull. Inst. Franc. Afr. Noire 16,
Sér. A: 795 (1954).
Adenopus breviflorus Benth.—Exell, Cat.: 184.

ANNOBON : Near Ambo, 24 m., pathside, 12 July 1959, Wrigley 19. Near
Ambo, 30 m. (common in other villages), hedges, roofs, etc., climber, 9 Aug. 1959,
Wrigley 254.

New record for Annobon.

Jeffrey (Kew Bull. 15 : 355 (1962)), in a new classification of the family Cucurbit-
aceae, includes Adenopus Benth. within the genus Lagenarza Ser.

LAGENARIA SICERARIA (Molina) Standl. in Publ. Field Mus. Nat. Hist., Bot. Ser. 3 :
435 (1930).

Cucurbita lagenaria L., Sp. Pl. 2: 1010 (1753).
Cucurbita siceraria Molina, Sagg. Stor. Nat. Chil.: 133, 355 (1782).

ANNOBON : Ambo village, 9 m., cultivated ; “ fruit used as calabash”’ ; 20 Aug.
1959, Melville 302.

New record for Annobon. Widespread and frequently cultivated in the tropics.
Calabash, Bottle-gourd or White Pumpkin.

LUFFA AEGYPTIACA Mill.— Exell, Cat.: 184.

ANNOBON : Ambo, near the expedition’s houses, 24 m., climbing over top branches
of Jatropha curcas, 12 July 1959, Melville 133. San Pedro, 15 m., village rubbish
heap, 2 Aug. 1959, Wrigley 229.

New record for Annobon.

[DIPLOCYCLOs PALMATUS (L.) Jeffrey in Kew Bull. 15 : 352 (1962).

Bryonia palmata L., Sp. Pl. 2: 1012 (1753).
Bryonopsis laciniosa Naud. in Ann. Sci. Nat., Sér. 4, Bot. 12: 141 (1859) pro parte.—
Exell, Cat.: 185.

Jeffrey (tom. cit.: 352-354) has adjusted the nomenclature of this widely distri-
buted palaeotropical species, which is recorded for S. Tomé. He adopts the generic
name Dzplocyclos (Endl.) Post & Kuntze.]

[ZEHNERIA GILLETII (De Wild.) Jeffrey in Kew Bull. 15 : 366 (1962).

Melothria cordifolia Hook. f. in Oliv., Fl. Trop. Afr. 2 : 563 (1871) ; non Zehneria cordifolia
Schweinf. ex Broun & Massey (1929).

Melothria gilletit De Wild. in Ann. Mus. Congo, Bot. Sér. 5, 3 : 140, t. 13 figs. 4-6 (1907).

Melothria capillacea sensu Exell, Cat.: 185 ; Suppl.: 22.

Occurs in S. Tomé and on the mainland of western tropical Africa from Liberia
to Gabon and Congo.

This and the following species belong to the palaeotropical genus Zehneria Endl.
which is separated by Jeffrey (tom. cit.: 343) from the exclusively New World
genus Melothria L.|

102 ANGIOSPERMS OF ANNOBON ISLAND

[ZEHNERIA SCABRA (L. f.) Sond. in Harv. & Sond., Fl. Cap. 2: 486 (1862).—Jeffrey
in Kew Bull. 15 : 369 (1962).
Bryonia scabra L. f., Suppl. Pl.: 423 (1781).
Zehneria scrobiculata Hochst. ex A. Rich., Tent. Fl. Abyss. 1 : 287 (1848).
Melothria minutifiora sensu Exell, Cat.: 185.
Recorded from S. Tomé ; widely distributed in the Old World tropics, extending
into South Africa. ]

ARALIACEAE
POLYSCIAS GUILFOYLEI (Bull) L. H. Bailey in Rhodora 18 : 153 (1916).
Aralia guilfoylei Bull, Cat. 1873 : 4 (1873).
Var. LACINIATA L. H. Bailey, loc. cit. (1916).
ANNOBON : Capelle San Juan, 4:5 m., garden of Capelle, 14 Aug. 1959, Melville

281.
New record for the islands. Introduced ; a cultivated plant of Polynesian origin.

RUBIACEAE
IxXORA COCCINEA L.—Exell, Suppl.: 24.
ANNOBON: Ambo, near the Mission, 15 m., cultivated land, 2 Aug. 1959,
Melville 278.
New record for Annobon. Introduced.

CoFFEA LIBERICA Bull ex Hiern.—G. Taylor in Exell, Cat.: 208.

ANNOBON : Ridge west of Crater Lake, 360 m., open woodland, 18 July 1959,
Wrigley 50.

New record for Annobon. Introduced.

BERTIERA ANNOBONENSIS G. Taylor.—G. Taylor in Exell, Cat.: 208. (Plate 8.)
ANNOBON : Near highest point of Crater, 480 m., forest ; ‘“‘ undershrub, two
specimens with white corolla, others fruit only ’’; 26 July 1959, Wrigley 93.
Endemic to Annobon. Wrigley 93 is topotypical material of Mildbraed 6502
(Herb. Berlin), one of the destroyed syntypes. The other syntype, Mildbraed 6760,
came from Santa Mina.

UNCARIA AFRICANA G. Don, Gen. Syst. 3 : 471 (1834).

ANNOBON : Pico Surcado and Monte Abecin, 570 m., forest climber, 11 Aug.
1959, Wrigley 304.

New record of the genus Uncaria Schreb. for the islands. This is the typical
variety of U. africana, distributed on the mainland of West Africa from Portuguese
Guinea eastwards and extending to Sudan, Uganda and Tanganyika.

[GEOPHILA NEURODICTYON (K. Schum.) Hepper in Kew Bull. 16 : 331 (1962).
Psychotria neurodictyon K. Schum.—G. Taylor in Exell, Cat.: 213.
Recorded from Principe.

ANGIOSPERMS OF ANNOBON ISLAND 103

The generic name Geophila D. Don (non Bergeret) has been proposed for conserva-
tion over Carinta W. F. Wight, the name adopted by G. Taylor (in Exell, Suppl.: 25)
for the three species previously listed by him (in Exell, Cat.: 217) under Geophila ;
cf. Taxon 9 : 88 (1960).]

OLDENLANDIA HERBACEA (L.) Roxb., Hort. Bengal.: 11 (1814).
Hedyotis herbacea L., Sp. Pl. 1 : 102 (1753).
ANNOBON : N.W. of the Island, 15 m., cassava plantation, 11 July 1959, Wrigley
Fi;
New record for the islands. Widely distributed in the Old World tropics.

COMPOSITAE
ADENOSTEMMA PERROTTETII DC.—Exell, Cat.: 222 ; Suppl.: 26.
ANNOBON : South of Crater Lake, near higher point of Crater, 420 m., cultivated
land, cassava plantations in the hills, 1 Aug. 1959, Melville 2z0.
New record for Annobon.

MIKANIA CORDATA (Burm. f.) B. L. Robinson.—Cannon in Exell, Suppl.: 27.

Mikania scandens sensu Exell, Cat.: 222.
Mikania sp., Cannon in Exell, Suppl.: 27.

ANNOBON : Near Ambo, 30 m., scrub, climber, 12 July 1959, Wrigley 27.
Previously recorded from Annobon as Mtkania sp.

(DICHROCEPHALA INTEGRIFOLIA (L. f.) Kuntze, Revis. Gen. Pl. 1 : 333 (1891).
Hippia integrifolia L. f., Suppl. Pl.: 389 (1781).
Dichrocephala bicolor (Roth) Schlecht.—Exell, Cat.: 223 ; Suppl.: 31.
The epithet integrifolia, though inapt, is the correct one for this widespread
species which is recorded for S. Tomé. ]

SYNEDRELLA NODIFLORA (L.) Gaertn.—Exell, Cat.: 226 ; Suppl.: 31.

ANNOBON:: North shore of Crater Lake, 265 m., pathside, 24 July 1959, Wrigley
80.

New record for Annobon.

ELEUTHERANTHERA RUDERALIS (Swartz) Schultz Bip. in Bot. Zeit. 24 : 164 (1866).
Melampodium ruderale Swartz, Fl. Ind. Occ. 3 : 1372 (1806).
ANNOBON : Near Capelle San Juan, 30 m., wayside in cultivated land, 10 Aug.
1959, Melville 257.
New record of the genus Eleutheranthera Poit. ex Bosc for the islands. E.
ruderalis is a pantropical weed.

CAMPANULACEAE
CEPHALOSTIGMA PERROTTETII A. DC., Monogr. Campan.: 118 (1830).
ANNOBON : Pico do Fogo, 360 m., among rocks, 24 July 1959, Wrigley go.
New record of the genus Cephalostigma A. DC. for the islands. C. perrotteti
is widespread in tropical Africa and tropical South America.

104 ANGIOSPERMS OF ANNOBON ISLAND

APOCYNACEAE
ALLAMANDA CATHARTICA L., Mant. Pl. Alt.: 214 (1771).
ANNOBON : Ambo, near the Mission, 15 m., 2 Aug. 1959, Melville 220.
New record of the genus Allamanda L. for the islands. A. cathartica, native of
tropical America, is widely cultivated in the tropics.

CATHARANTHUS ROSEUS (L.) G. Don.—Philipson in Exell, Cat.: 239.
ANNOBON : Ambo, 27 m., waste ground, 19 July 1959, Wrigley 58.
New record for Annobon. Introduced.

PLUMERIA RUBRA L.—Philipson in Exell, Cat.: 241.
ANNOBON : Near Ambo, 24 m., in front of church, planted, 18 Aug. 1959, Wrigley

293.
New record for Annobon. Introduced.

ASCLEPIADACEAE

ASCLEPIAS CURASSAVICA L.—Exell, Cat.: 242 ; Suppl.: 34.

ANNOBON : Ambo, 180 m. S.W. of church, 22 m., cleared ground, 10 July 1959,
Melville ro2.

New record for Annobon. Introduced.

LOGANIACEAE
(By J. Lewis)

NUXIA CONGESTA R. Br. ex Fresen. in Flora 21 (2) : 606 (1838).—Bruce & Lewis in

Five iiop: Air, Logan; 44, fig’ §.(7, 8) (1900):

Nuxia congesta R. Br. in Salt, Voy. Abyss., App.: 63 (1814), nom. nud.

Var. CONGESTA. (Plate 9.)

Nuxia angolensis Gilg in Notizbl. K. Bot. Gart. & Mus. Berl. 1 : 74 (1895).

Nuxia mannii Gilg in Engler, Bot. Jahrb. 30 : 376 (1901).

Nuxia annobonensis Mildbr. in Wiss. Ergebn. Zweit. Deutsch. Z.-Afr.-Exped. 2: 163

(1922), nom. nud.
Lachnopylis mannii (Gilg) Hutch. & Moss in Hutch. & Dalziel, Fl. W. Trop. Afr. 2:

20, fig. 185 (1931).
Lachnopylis annobonensis Mildbr.—Philipson in Exell, Cat.: 244.
Lachnopylis angolensis (Gilg) Philipson in Exell, Cat.: 245 (1944).

ANNOBON : Pico do Fogo, 2 m. and less below top of Pico, rocky mountainside,
12 Aug. 1959, Melville 277.

Widely distributed on the mountains of tropical Africa and extending into South
Africa ; previously recorded for Annobon under the name Lachnopylis annobonensis,
of which Melville 277 is an exact topotype. The holotype of L. annobonensis,
Miuldbraed 6561 (Herb. Berlin), is destroyed, but there is an isotype in the British
Museum Herbarium accompanied by a photograph of the holotype which is
reproduced in Plate g.

The opinions expressed by Bruce and Lewis (loc. cit.) and by Verdoorn (Bothalia
7 : 14 (1958)) concerning the great variability and wide range of this species are

ANGIOSPERMS OF ANNOBON ISLAND 105

confirmed by more recent workers in Floras concerning West and South Africa
about to be published, and I have now no hesitation in including N. angolensis and
N. mannii as well as Lachnopylis annobonensis. The Annobon plant is very clearly
the common form of the species which has been known as N. angolensis on the
neighbouring mainland. On S. Tomé, however, the species is represented by a
plant which may be distinguished varietally.

[Var. thomensis (Philipson) J. Lewis, stat. nov.

Lachnopylis thomensis Philipson in Exell, Cat.: 245 (1944).—Exell, Addit.: 462.

Known from S. Tomé and Mount Kilimanjaro in Tanganyika.

Specimens collected by Monod on the Pico in S. Tomé (Monod 11954, 11989) have
unusually large elliptic leaves, about 15 cm. long and 6 cm. broad, bearing distinct
traces of a white dendroid indumentum along the midrib and nerves beneath.
However, they are exactly matched by a specimen collected by Haarer at a compar-
able altitude on Mount Kilimanjaro (Herb. Kew) which has always been accepted
as N.congesta. The occurrence of N. congesta at similar altitudes on other mountains
makes subspecific rank unsuitable for these variants, but their moderate distinctness
justifies recognition at the varietal level. |

GENTIANACEAE
CANSCORA DECUSSATA (Roxb.) J. A. & J. H. Schult., Mant. 3 : 229 (1827).

Pladeva decussata Roxb., Hort. Bengal.: 10 (1814), nom. nud.
Pladera decussata Roxb., Fl. Ind., ed. Carey, 1 : 418 (1820).
ANNOBON: N.W. of the Island, 15 m., pathside, 11 July 1959, Melville 137.
New record of the genus Canscora Lam. for the islands. C. decussata is widespread
in the tropics of the Old World.

EXACUM QUINQUENERVIUM Griseb., Gen. & Sp. Gentian.: 112 (1839).
ANNOBON : N.W. of the Island, 15 m., cassava plantation, 11 July 1959, Wrigley
9.
New record of the genus Exacum L. for the islands. E. guinquenervium is widely

distributed in tropical Africa, Madagascar and the Mascarene Islands.

CONVOLVULACEAE
IPOMOEA ALBA L., Sp. Pl. 1 : 161 (1753).
Calonyction aculeatum (L.) House.—Exell, Cat.: 249.

ANNOBON : South of Crater Lake, 450 m., opening in forest, climbing on plants,
1 Aug. 1959, Melville 207.

New record for Annobon.

Calonyction Choisy is now included in Ipomoea L., and C. aculeatum takes the
name I. alba.

IPOMOEA INVOLUCRATA Beauv., Fl. Oware & Benin 2: 52, t. 89 (1816).

ANNOBON:: North of San Pedro, 420 m., clearings in forest, 21 Aug. 1959,
Melville 305.

New record for the islands. Widespread in tropical Africa.

106 ANGIOSPERMS OF ANNOBON ISLAND

IPOMOEA MAURITIANA Jacq.—Exell, Addit.: 462.
Ipomoea digitata sensu Exell, Cat.: 250.
ANNOBON : Crater Lake, 265 m., lake edge, on Ficus and Elaeis, climber, 31 July

1959, Melville 206.
New record for Annobon.

IPpoMOEA NIL (L.) Roth, Catalect. Bot. 1 : 36 (1797).
Convolvulus nil L., Sp. Pl., ed. 2, 1 : 219 (1762).
ANNOBON : South of the Administrator’s house and just behind the expedition’s
houses, 24 m., shady pathside, running over ground, 10 July 1959, Melville 103.
Newrecord for theislands. Introduced ; a widespread tropical species of American
origin.

SOLANACEAE
(By W. T. Stearn)
LYCOPERSICON ESCULENTUM Mill.—Exell, Cat.: 252.
Var. CERASIFORME (Dunal) Alef.—Exell, Cat.: 252.
ANNOBON : San Pedro, 3 m., sea shore, cultivated, 11 Aug. 1959, Melville 262.
New record for Annobon. Introduced.

SOLANUM DASYPHYLLUM Schumach. in K. Danske Vid. Selsk. Naturvid. & Math.
Afhandl. 3 : 146 (1828).—Bitter in Fedde, Repert. Sp. Nov., Beih. 16: 188
(1923).

Solanum duplosinuatum Klotzsch.—Exell, Cat.: 253.
ANNOBON : Between Pico do Fogo and San Pedro, 450 m., banana plantation,

26 July 1959, Wrigley 95.

New record for Annobon ; recorded (with doubt) for Principe under the name

S. duplosinuatum. Widespread in tropical Africa, extending into Natal.

SOLANUM NODIFLORUM Jacq., Collect. 2 : 288 (1788).
Solanum nigrum sensu Exell, Cat.: 253 ; Suppl.: 36.

ANNOBON : Pico do Fogo, S.W. side, 390 m., bare soil between rocks, recently
burnt, 25 July 1959, Melville 188.

Recorded for Principe, S. Tomé and Annobon under the name S. nigrum. Wide-
spread in the tropics, where it has been much confused with the closely allied
S. nigrum L.

PHYSALIS ANGULATA L.—Exell, Cat.: 254.

ANNOBON: N.W. of the Island, 15 m., cassava plantation, 11 July 1959,
Wrigley 124.

New record for Annobon. Probably introduced.

PHYSALIS MICRANTHA Link, Enum. Pl. Hort. Bot. Berol. 1: 181 (1821).—O. E.
Schulz in Urban, Symb. Antill. 6 : 147 (1909).

ANGIOSPERMS OF ANNOBON ISLAND 107

Physalis minima sensu C. H. Wright in Dyer, Fl. Trop. Afr. 4 (2) : 247 (1906).

ANNOBON : N.W. of the Island, 15 m., cassava plantation, 11 July 1959, Melville
125. Pico do Fogo, 390 m., bare soil, 25 July 1959, Melville 195.

New record for the islands. Widespread in tropical Africa and also recorded for
India and tropical America.

CAPSICUM ANNUUM L.—Exell, Cat.: 254.

ANNOBON : Valley between Pico do Fogo and San Pedro, 450 m., banana planta-
tion, 26 July 1959, Wrigley 94, 94a. Highest point of Crater, 420 m., cultivated
land, 1 Aug. 1959, Melville 221. South of Santa Cruz, cultivated land, 19 Aug.
1959, Melville 314.

New record for Annobon. Cultivated.

DATURA CANDIDA (Pers.) Safford.—Exell, Suppl.: 36.
ANNOBON : Ambo, in front of church, 15 m., garden, 22 Aug. 1959, Melville 312.
New record for Annobon. Introduced ; widely spread as a cultivated plant in
tropical and subtropical countries.

NICOTIANA TABACUM L.—Exell, Cat.: 256.
ANNOBON : Ambo, 27 m., waste ground ; “not (obviously) cultivated ; not

used’; 19 July 1959, Wrigley 59.
New record for Annobon. Introduced.

SCROPHULARIACEAE
LINDERNIA DIFFUSA (L.) Wettst.—Exell, Cat.: 256.
Var. DIFFUSA.

ANNOBON:: N.W. of the Island, 15 m., cassava plantation, 11 July 1959, Wrigley
IO.

New record for Annobon.

Var. PEDUNCULATA (Benth.) Skan.—Exell, Cat.: 256 ; Suppl.: 36.
ANNOBON : Top of Pico Surcado, 11 July 1959, Wrigley roa.
New record for Annobon.

ALECTRA SESSILIFLORA (Vahl) Kuntze, Revis. Gen. Pl. 2 : 458 (1891).
Gerardia sessiliflora Vahl, Symb. Bot. 3 : 79 (1794).
Var. MONTICOLA (Engler) Melch. in Notizbl. Bot. Gart. & Mus. Berl.-Dahl. 15 :
126 (1940).—Hepper in Kew Bull. 14 : 406 (1960).
Melasma indicum var. monticola Engler, Bot. Jahrb. 30 : 402 (1901).
Alectra communis Hemsl.—Exell, Cat.: 257.
ANNOBON : Pico do Fogo, S.W. side, 390 m., burnt area, bare ground, 25 July
1959, Melville 189.
New record for Annobon. Recorded for Principe ; widespread in tropical

Africa, Madagascar, Mauritius and south-eastern Asia from India to Formosa and
the Philippines.

108 ANGIOSPERMS OF ANNOBON ISLAND

ACANTHACEAE
PHAULOPSIS MICRANTHA (Benth.) C. B. Clarke.—Exell, Cat.: 261.
ANNOBON : N.W. of the Island, 15 m., cassava plantation, 11 July 1959,
Melville 120.
New record for Annobon.

DICLIPTERA VERTICILLATA (Forsk.) C. Christens. in Dansk Bot. Ark. 4 (3) : 11 (1922).

Dianthera verticillata Forsk., Fl. Aegypt.-arab.: cClll, 9 (1775).
Justicia umbellata Vahl, Enum. Pl. 1: 111 (1805).
Dicliptera umbellata (Vahl) Juss.—Exell, Cat.: 263 ; Addit.: 463.

ANNOBON : Near Ambo and near Crater Lake, near streams, etc., 18 July 1959,
Wrigley 49.

New record for Annobon. The species occurs in Principe and S. Tomé and is
widespread in tropical Africa, extending into Arabia and India.

[VERBENACEAE]
[AVICENNIA GERMINANS (L.) L., Sp. Pl., ed. 3, 2 : 891 (1764).—Stearn in Kew Bull.
13: 34 (1958).—Exell, Aditam.: 86; Addit.: 464.
Avicennia africana Beauv.—Exell, Cat.: 265.
The earliest publication of the combination A. germinans was in 1764 as given
above. The species is recorded from S. Tomé ; it occurs on the coasts of tropical
America as well as West Africa. ]

-

LABIATAE
LEONOTIS NEPETIFOLIA (L.) Ait. f—G. Taylor in Exell, Cat.: 266.—Exell, Suppl.: 38.
ANNOBON : Ambo, by the expedition’s houses, 15 m., waste ground and common
there ; ‘‘ natives dry and hang up to keep off mosquitoes ’’; 13 July 1959, Melville
I4t.
New record for Annobon.

OcIMUM BASILIcUM L.—G. Taylor in Exell, Cat.: 266.
ANNOBON : Ambo village, 15 m., pathside, 15 July 1959, Wrigley 4o.
New record for Annobon.

NYCTAGINACEAE
BoOERHAVIA COCCINEA Mill., Gard. Dict., ed. 8, n. 4 (1768).—Meikle in Hutch. &
Dalziel, Fl. W. Trop. Afr., ed. 2,1: 178, fig. 66 (1954).
Boerhavia diffusa sensu Exell, Cat.: 268 pro parte.
PRINCIPE : Without locality, Sept. 1853, Welwitsch 5391.
S. Tomé : Without locality, Don. Rio do Ouro, 240 m., July 1885, Moller 590.
Blu-blu, 130 m., 26 Sept. 1912, Watt 7090. Roca Rozema, 13 Oct. 1912, Watt

7350, 7357:
ANNOBON : Without locality, 15 Feb. 1933, Exell 901. San Antonio, 60 m.,

village, between houses, 8 Aug. 1959, Melville 252.

ANGIOSPERMS OF ANNOBON ISLAND 109

New records for Principe, S. Tomé and Annobon. Native of tropical America,
now widespread in the tropics of both America and Africa.

Since the publication of my Catalogue in 1944 Meikle (loc. cit.) has distinguished
two species within the West African material referred to B. diffusa L. Re-examin-
ation of the specimens in the British Museum Herbarium cited in the Catalogue
shows that they all belong to B. coccinea as defined by Meikle.

BOERHAVIA DIFFUSA L.—Exell, Cat.: 268 excl. parte.

ANNOBON : Lower region, 0-150 m., 15 Feb. 1933, Exell 880. Towards Pico do
Fogo, south of Ambo, go m., pathside in open semi-cultivated country, 10 July
1959, Melville 106.

Recorded for Annobon by Mildbraed, but the identity of his plant is now uncertain.
The record for Principe, and at least some of the records for S. Tomé, belong to
B. coccinea as indicated above under that species.

AMARANTHACEAE
AMARANTHUS SPINOSUS L.—Exell, Cat.: 270.
ANNOBON : San Pedro, 3 m., pathside in village, 21 July 1959, Melville 173.
New record for Annobon.

AMARANTHUS VIRIDIS L.—Exell, Cat.: 270.

ANNOBON : N.W. corner of Crater Lake, 265 m., dry stream bed, 24 July 1959,
Melville 179.

New record for Annobon.

ACHYRANTHES ASPERA L.—Exell, Cat.: 270.
ANNOBON : Crater Lake, 270 m., forest paths, 25 July 1959, Melville 192.
New record for Annobon.

CYATHULA PROSTRATA (L.) Blume.—F xell, Cat.: 271.

ANNOBON : Path to Pico Surcado (Monte Santa Mina), 510 m., dense vegetation
in mist-forest, 21 July 1959, Melville 174.

New record for Annobon.

ALTERNANTHERA MARITIMA (Mart.) St.-Hil., Voy. Brésil 2: 437 (1833).—Keay in
Hutch. & Dalziel, Fl. W. Trop. Afr., ed. 2, 1 : 154 (1954).
Bucholzia maritima Mart., Nov. Gen. & Sp. Pl. 2: 50, t. 147 (1826).
Telanthera maritima (Mart.) Moq.—Exell, Cat.: 271 ; Aditam.: 86.
ANNOBON : Near Bird Island, N.W. coast, 6 m., sea cliff with wind-blown spray,
17 Aug. 1959, Wrigley 284.
Recorded for Principe, S. Tomé and Annobon.

Keay and other recent authors include Telanthera R. Br. under Alternanthera
Forsk.

ALTERNANTHERA SESSILIS (L.) R. Br. ex DC., Cat. Pl. Hort. Bot. Monspel.: 4, 77
(1813).—Exell, Cat.: 272 ; Suppl.: 30.

110 ANGIOSPERMS OF ANNOBON ISLAND

ANNOBON : Dry stream east of Ambo, 30 m., between rocks, 1 Aug. 1959, Wrigley
205. Near Bird Island, N.W. coast, 9 m., sea cliffs, sometimes in wind-blown
spray, 17 Aug. 1959, Melville 283.

The earliest publication of the combination A. sessilis appears to be that given
above. The species is recorded for Principe, S. Tomé and Annobon.

BASELLACEAE
BASELLA ALBA L.—Fxell, Cat.: 274.
ANNOBON : San Pedro and Ambo, 15 m., roof of houses, vine rooted in the
ground ; “ used as pot-herb’”’; 21 July 1959, Wrigley 67.
New record for Annobon. Probably introduced.

PIPERACEAE
(By W. T. Stearn)

PEPEROMIA HYGROPHILA Engler.—Exell, Cat.: 277.
Peperomia annobonensis Mildbr.—Exell, Cat.: 276.

ANNOBON: Pico Surcado, 510 m., mist-forest, very abundant epiphyte, 21
July 1959, Wrigley 7o.

P. hygrophila was based by Engler on specimens collected at 2,500-2,600 m.
altitude on Cameroons Mountain, evidently in a moist habitat as the epithet implies ;
an isosyntype (Preuss 805) in the British Museum Herbarium represents a robust
luxuriant individual with cuneate-based leaf-laminas 1-5-4 cm. broad and a flowering
spike about 8 cm. long, these somewhat exceeding the dimensions given by Engler
in his original description. Exell 156 from virgin forest on S. Tomé approaches
Preuss’s plant in luxuriance, having laminas I-35 cm. broad, some cuneate, others
almost rounded at the base, but with spikes 2-5-4:5 cm. long. Other specimens
from S. Tomé bridge the gap between this and a small-leaved plant from the same
island (Monod 11744) with laminas rounded or almost subcordate at the base and
0:5-1°4 cm. broad. In Monod 12228 the lamina varies from almost circular with
rounded base to narrowly obovate with narrowly cuneate base. The spike in
S. Tomé material varies in length from about 8 cm. to 2cm. I have examined the
holotype of P. annobonensis (Mildbraed 6532) which is preserved at Berlin and which
was collected on Annobon in dry Steganthus-Elaeis forest at 250-350 m. altitude.
It differs from Wrigley’s mist-forest material in having mostly elliptic rather than
mostly circular laminas, but both these Annobon plants come well within the range
of variation of P. hygrophila as manifested on S. Tomé and Cameroons Mountain.

LAURACEAE
PERSEA AMERICANA Mill.—Exell, Cat.: 280.
ANNOBON : S.W. of Crater Lake, 390 m., open forest, 5 Aug. 1959, Melville 239.
New record for Annobon. Introduced. Avocado.

LORANTHACEAE
(By W. T. Stearn)

VISCUM ENGLERI Tiegh. in Bull. Soc. Bot. France 43: 190 (1896).—Sprague in
Dyer, Fl. Trop. Afr. 6 (1) : 405 (1911).

ANGIOSPERMS OF ANNOBON ISLAND Il

ANNOBON : Edge of forest near Pico do Fogo, 330 m., parasite on Fucus
clarencensis, 24 July 1959, Wrigley 86.

New record for the islands. Also in Angola and Tanganyika.

The specimen, which has only unripe fruit, belongs to the small group of African
species of Viscum with conspicuously flattened stems and the leaves reduced to
scales. In the key to the tropical African species given by Sprague (tom. cit.:
394-395) it runs down to V. engleri, hitherto recorded only from Tanganyika ;
and it comes within the range of variation of this species as indicated by specimens
from the Usambara Mountains (the locus classicus) and the Sagara Mountains.
Specimens of what is evidently the same species collected by Gossweiler (n. 9884,
Herb. Kew) in the Cuanza Sul region of Angola suggest that V. englert may be
widespread but sporadic in tropical Africa.

EUPHORBIACEAE

EUPHORBIA PULCHERRIMA Willd. ex Klotzsch in Allgem. Gartenzeit. 2 : 27 (1834).
ANNOBON : Santa Cruz, 30 m. (also Ambo), planted by chapel, 8 Aug. 1959,
Melville 269.
New record for the islands. Introduced ; native of Mexico and Central America.
Poinsettia.

[CLEISTANTHUS LIBERICUS N. E. Br.—Exell, Cat.: 286.

In the Catalogue I listed this species from S. Tomé, but with a query. Dr. J.
Léonard considers (and I agree) that the S. Tomé plant was correctly identified as
C. libericus by Hutchinson. ]

THECACORIS ANNOBONAE Pax & Hoffm.—Exell, Cat.: 287. (Plate ro.)

ANNOBON : Pico Surcado, 420 m., forest ; “‘shiny leaves ; very red wood ;
racemes of flowers with 5 stamens”; 21 July 1959, Wrigley 66. Near highest
point of Crater, 480 m., forest, 26 July 1959, Wrigley 92.

Endemic to Annobon. This identification is probably correct but there is a slight
element of doubt as the species was described from female inflorescences and fruit
while Wrigley collected male inflorescences and fruit, so that his material can be
compared only partly with the original description. JT. annobonae was described
as having “ capsulae cocci pilosi’’; the fruits of Wrigley g2 are nearly glabrous but
have vestiges of anindumentum. Although there is a suggestion that there may be
another species of Thecacorts on the island (Exell, Cat.: 288) Wrigley’s specimens
nevertheless in all probability represent the species (T. annobonae) collected by
Mildbraed on the rim of the North Crater at c. 500 m. altitude and said to be very
common.

PHYLLANTHUS RETICULATUS Poir. in Encycl. Méth., Bot. 5 : 298 (1804).—G. L.
Webster in Journ. Arnold Arb. 38 : 57, fig. 7 (1957).
ANNOBON : East of Quioveo (Pico del Centro), edge of clearing in forest, straggler,
1 Aug. 1959, Melville 212.
New record for the islands. Widespread in the Old World tropics ; introduced
into the West Indies.

112 ANGIOSPERMS OF ANNOBON ISLAND

JATROPHA MULTIFIDA L.—Exell, Cat.: 293 ; Suppl.: 42.

ANNOBON : Ambo, near the Post Office, 12 m., pathside, cultivated patch,
16 July 1959, Melville 156.

New record for Annobon. Introduced.

CODIAEUM VARIEGATUM (L.) Blume.—Exell, Cat.: 296.

ANNOBON : Due south of Crater Lake, 420 m., forest near path to Quioveo,
1 Aug. 1959, Melville 208.

New record for Annobon. Introduced.

Discoclaoxylon pubescens (Pax & Hoffm.) Exell, stat. nov. (Plate 11.)
Discoclaoxylon occidentale var. pubescens Pax & Hoffm. in Engler, Bot. Jahrb. 58, Beibl.
130 : 39 (1923).—Exell, Cat.: 299.

ANNOBON : Highest point of Crater, 480 m., forest ; “‘ large leaves, 2/5 phyllo-
taxis ; catkin-like inflorescence ; 4-merous green flowers ’’; 26 July 1959, Wrigley
96.

Endemic to Annobon. There is little doubt that Wrigley’s plant is identical
with D. occidentale var. pubescens, described from rather insufficient female material
consisting of three syntypes : Muldbraed 6492, 6555 and 6751 (Herb. Berlin, des-
troyed). The male flowers in Wrigley’s material are considerably larger than those
of D. occidentale (Muell. Arg.) Pax & Hoffm., measuring about 8 mm. in diameter
when expanded. There are about five male flowers to each bract. They seem to
expand one at a time and the pedicel is then 5-8 mm. long. There are 10-12
stamens. The leaves are similar to those of D. occidentale but are more hairy on
the under surface, the petiole and the base of the midrib being tomentellous. I
consider that these differences justify separate specific rank for the Annobon plant,
and that D. occidentale is confined to S. Tomé.

Discoclaoxylon (originally Claoxylon sect. Discoclaoxylon Muell. Arg.) consists
of a group of West African species geographically separated from Claoxylon A. Juss.
and sufficiently distinct to warrant the generic rank given them by Pax and Hoffmann
(in Engler, Pflanzenr. 4 (147, 7) : 137 (1914)). Hutchinson and Dalziel (Fl. W.
Trop. Afr. 1 : 301 (1928)) reunited Discoclaoxylon with Claoxylon and this treatment
was followed by Keay (in Hutch. & Dalziel, op. cit., ed. 2, 1: 401 (1958)). The
main differences between the two genera, according to Pax and Hoffmann (tom.
cit.: 76, 100, 137), are :

Claoxylon : extrastaminal disk none, but small glands usually present between

the stamens ; stamens 10-200 or more (very rarely fewer than 15).

Discoclaoxylon : extrastaminal disk urceolar, entire or lobulate ; stamens 6-12.
Claoxylon does not occur on the continent of Africa but is distributed from
Madagascar and the Mascarene Islands eastwards through southern Asia to Australia
and Polynesia.

The genus Discoclaoxylon now comprises four species : D. pedicellare (Muell. Arg.)
Pax & Hoffm. (Fernando Po), D. occidentale (Muell. Arg.) Pax & Hoffm. (S: Tomé),
D. pubescens (Pax & Hoffm.) Exell (Annobon) and D. hexandrum (Muell. Arg.) Pax

ANGIOSPERMS OF ANNOBON ISLAND 113

& Hoffm. (Fernando Po, Cameroons and Congo). The last-named species, which
has (3-) 6-8 stamens (the others have 10-12) is the only one so far found on the
mainland of Africa. When male and female flowers and fruits of all four are known
their relationship may become clear, but it is already apparent that the distribution
is ‘“‘normal’’ with one West African species reaching Fernando Po, one endemic
on that island, one on S. Tomé and a fourth on Annobon. The genus has not been
recorded for Principe. It is one of the few “ island genera ’”’ with more species on
the islands than on the mainland (cf. Calvoa in the Melastomataceae which has
about half its known species on the islands).

It seems desirable to typify the generic name Discoclaoxylon. Claoxylon sect.
Discoclaoxylon Muell. Arg. (Flora 47 : 437 (1864)) was founded on three species,
C. pedicellare Muell. Arg., C. occidentale Muell. Arg. and C. hexandrum Muell. Arg.,
without indication of type ; all three species were known only from male material.
The position was unchanged when the section was later reviewed by its author (in
DC., Prodr. 15 (2) : 779 (1866)). When Pax and Hoffmann raised the section to
generic rank as Discoclaoxylon the third species (D. hexandrum) had become known
from the mainland of West Africa and female material had become available, so
that quite a large part of their generic description (relating to ovary, capsule and
seeds) came from this species, female material being still unknown in the other two
species. It is also quite possible that D. pedicellare and D. occidentale are now
extinct and complete material may never become available. Taking these points
into consideration it seems best to select D. hexandrum as type of Discoclaoxylon.

ACALYPHA ANNOBONAE Pax & Hoffm.—Exell, Cat.: 299. (Plate 12.)
ANNOBON : Ridge west of Crater Lake, 360 m., open woodland ; “‘ female and
male flowers on same shrub ; low shrub 1-2 m.”’; 18 July 1959, Wrigley 55.
Endemic to Annobon. Wrigley’s plant is practically a topotype. The holotype,
Miuldbraed 6538 (Herb. Berlin), has been destroyed.

RICINUS COMMUNIS L.—Exell, Cat.: 301.
ANNOBON : Ambo village, 15 m., pathside, 15 July 1959, Wrigley 4r.
New record for Annobon. Probably introduced.

MORACEAE

ARTOCARPUS COMMUNIS J. R. & G. Forst.—G. Taylor in Exell, Cat.: 304.
ANNOBON: Near Ambo, 6 m., waste ground near sea, not seen to regenerate,
18 Aug. 1959, Wrigley 292.
New record for Annobon. Introduced. Breadfruit.

ARTOCARPUS HETEROPHYLLUS Lam. in Encycl. Méth., Bot. 3 : 209 (1789).—Jarrett
in Journ. Arnold Arb. 40 : 334 (1959).

Artocarpus integer sensu G. Taylor in Exell, Cat.: 304.

ANNOBON:: Near highest point of Crater, 480 m. (also one sterile specimen on
shore of Lake), forest, edge of clearing, 26 July 1959, Wrigley gr.

114 ANGIOSPERMS OF ANNOBON ISLAND

New record for Annobon ; recorded for Principe and S. Tomé under the name
A. integer. Introduced. Jack Fruit.

The name A. integer (Thunb.) Merr. applies to a different species (cf. Jarrett,
tom. cit.: 329).

URTICACEAE
URERA CAMEROONENSIS Wedd.—G. Taylor in Exell, Cat.: 312.
ANNOBON : Common all over the Island in forest, forest scrambler, often forming
lianes, 31 July 1959, Wrigley 223.
New record for Annobon.

ORCHIDACEAE
(By V. S. Summerhayes)

EPIPOGIUM ROSEUM (D. Don) Lindl. in Journ. Proc. Linn. Soc. Lond., Bot. I: 177
(1857).
Limodorum roseum D. Don, Prodr. Fl. Nepal.: 30 (1825).

ANNOBON : South of Crater Lake and east of Quioveo (Pico del Centro), 450 m.,
humus below trees with locally sparse fern flora on ground, 1 Aug. 1959, Melville 213
(Herb. Kew).

New record of the genus Epipogium R. Br. for the islands. EE. roseum is widely
but sporadically distributed in tropical Africa (Ghana, Cameroons, Fernando Po,
Congo, Angola, Uganda), Indo-Malaysia, Australia and New Hebrides.

PLATYLEPIS GLANDULOSA (Lindl.) Reichb. f—Dandy in Exell, Cat.: 319.—Exell,
Suppl 47,
ANNOBON : South of Santa Cruz (also Crater Lake, N.E. side), under forest,
24 July 1959, Melville 184 (Herb. Kew).
New record for Annobon. The species extends from tropical Africa into Natal.

CORYMBORKIS CORYMBOSA Thou., Hist. Pl. Orch., Tab. Esp. 1 (1822).—Summerh.
in Kew Bull. 11 : 224 (1956).
Corymborkis welwitschii (Reichb. f.) Kuntze.—Dandy in Exell, Cat.: 320.

ANNOBON : Between Crater Lake and Monte Abecin, 450 m., forest, 19 Aug.
1959, Melville 299 (Herb. Kew).

New record for Annobon. The species is recorded for S. Tomé and is generally
distributed in tropical Africa, extending into Natal and eastern Cape Province ;
it also occurs in Madagascar and Réunion.

LIPARIS WELWITSCHII Reichb. f. in Flora 48 : 184 (1865).

ANNOBON : Pico Surcado, 600 m., beneath canopy of ferns in forest, 11 Aug.
1959, Melville 266 (Herb. Kew).
New record for the islands. Also in Angola.

ANGIOSPERMS OF ANNOBON ISLAND 115

POLYSTACHYA RIDLEYI Rolfe.—Dandy in Exell, Cat.: 323.

ANNOBON : South of Crater Lake, 510 m., epiphytic on Coffea trees in particular,
14 July 1959, Melville 154 (Herb. Kew).

New record for Annobon. Hitherto believed to be endemic to S. Tomé.

BULBOPHYLLUM MELANORRHACHIS (Reichb. f.) Reichb. f. ex De Wild., Pl. Bequaert.
E 3) 93° (1921):
Megaclinium melanorrhachis Reichb. f. in Gard. Chron., Ser. 2, 4 : 162 (1875).
Bulbophyllum melanorrhachis Reichb. f., loc. cit. (1875), nom. syn.
ANNOBON : Ridge west of Crater Lake, 510 m., epiphyte, 14 July 1959, Wrigley
31 (Herb. Kew).
New record for the islands. Also on the mainland of West Africa from Guinea to
Nigeria, Gabon and Congo.

BOLUSIELLA TALBOTII (Rendle) Summerh. in Hutch. & Dalziel, Fl. W. Trop. Afr. 2 :
456 (1936).
Angraecum talbotit Rendle, Cat. Pl. Talbot : 108, t. 15 figs. 6, 7 (1913).
ANNOBON : Ridge west of Crater Lake, 510 m., epiphyte, 14 July 1959, Wrigley
33 (Herb. Kew).
New record of the genus Bolusiella Schlechter for the islands. B. talbotii occurs
on the mainland of West Africa from Sierra Leone to Nigeria.

AMARYLLIDACEAE

HYMENOCALLIs LITTORALIS (Jacq.) Salisb. in Trans. Hort. Soc. Lond. 1 : 338 (1812).
Pancratium littorale Jacq., Select. Stirp. Amer. Hist.: 99, t. 179 fig. 94 (1763).
ANNOBON : Mission, Ambo, 15 m., garden, 4 Aug. 1959, Melville 234.

New record of the genus Hymenocallis Salisb. for the islands. H. littoralis is
introduced into West Africa from tropical America.

DIOSCOREACEAE

DIOscOREA ALATA L.—Dandy in Exell, Cat.: 344.—Exell, Suppl.: 49.
ANNOBON : Ambo, path to San Pedro, 24 m., cassava plantation, 22 Aug. 1959,
Melville 308.

New record for Annobon. Introduced.

DIOSCOREA CAYENENSIS Lam.—Dandy in Exell, Cat.: 345.—Exell, Suppl.: 4g.
ANNOBON : Ambo, south of the Governor’s house, near footpath, 10 July 1959,
Wrigley 2. South of Santa Cruz, 420 m., openings for cultivation, common,
scrambling, 7 Aug. 1959, Wrigley 247.
New record for Annobon.

DIoscOREA DUMETORUM (Kunth) Pax.—Dandy in Exell, Cat.: 345.—Exell,
Suppl.: 49.

116 ANGIOSPERMS OF ANNOBON ISLAND

ANNOBON : South of Ambo, towards Pico do Fogo, 135 m., climber, 10 July
1959, Melville rrr. North of Santa Cruz, on coast path, 45 m., opening in forest,
climber, 8 Aug. 1959, Wrigley 253.

New record for Annobon.

LILIACEAE

CORDYLINE FRUTICOSA (L.) A. Chev., Jard. Bot. Saigon : 66 (1919).
Convallaria fruticosa L., Herb. Amboin.: 16 (1754).
ANNOBON : Santa Cruz and near Quioveo, 420 m., forest near path, 17 July

1959, Wrigley 48.
New record of the genus Cordyline Commers. ex Juss. for the islands. C. fruticosa

is a native of south-east Asia, introduced into tropical Africa and America.

SANSEVIERIA METALLICA Géréme & Labroy in Bull. Mus. Hist. Nat. Par. 9: 170,
173, fig. 2 (1903).
ANNOBON : Ambo, near the Mission, 15 m., cultivated ground, 22 Aug. 1959,
Wrigley 309.
New record for Annobon. Native of tropical Africa ; introduced into other
parts of the tropics. Species of Sansevieria (unnamed) have been introduced into
S. Tomé (cf. Dandy in Exell, Cat.: 347) but none is reported from Principe.

COMMELINACEAE

[STANFIELDIELLA IMPERFORATA (C. B. Clarke) Brenan in Kew Bull. 14 : 284 (1960).
Buforrestia imperforata C. B. Clarke.—Dandy in Exell, Cat.: 347.
The genus Stanfieldiella Brenan has recently been separated from Buforrestia
C. B. Clarke (cf. Brenan, tom. cit.: 280). S.imperforata var. imperforata is recorded
from Principe and S. Tomé.]

COMMELINA BENGHALENSIS L., Sp. Pl. 1: 41 (1753).

ANNOBON : San Pedro, 15 m., streamside, 2 Aug. 1959, Wrigley 228.

New record for the islands. Widely distributed in tropical and subtropical
regions of the Old World.

ARACEAE
XANTHOSOMA SAGITTIFOLIUM (L.) Schott in Schott & Endl., Melet. Bot.: 19 (1832).
Arum sagittifolium L., Sp. Pl. 2 : 966 (1753).
ANNOBON : Above Crater Lake, 360-480 m., open woodland, probably planted,

16 July 1959, Wrigley 44.
New record of the American genus Xanthosoma Schott for Annobon. X.
sagittifolium is introduced into tropical Africa from tropical America where it is

cultivated for its edible tubers.

ANGIOSPERMS OF ANNOBON ISLAND L177

CYPERACEAE
[HyPOLYTRUM. GRANDE (Uitt.) Koyama in Journ. Fac. Sci. Univ. Tokyo, Sect. 3,
8 : 68 (1961).
Principina grandis Uitt.—Dandy in Exell, Cat.: 355.
Koyama (loc. cit.); in his recent new arrangement of the family Cyperaceae,
has included Principina Uitt. in Hypolytrum Rich. Principina was previously
regarded as a monotypic genus endemic to Principe. |

CYPERUS PEDUNCULATUS (R. Br.) Kern in Act. Bot. Neer] 7 : 798, fig. 4 (1958).

Remirea maritima Aubl.; non Cyperus maritimus Poir.—Dandy in Exell, Cat.: 356.—
Exell, Aditam.: 87.
Remirea pedunculata R. Br., Prodr. Fl Nov. Holl.: 236 (1810),
Recorded for Principe and Annobon, but not collected by the Cambridge
Expedition.
Remirea Aubl. is included in the large genus Cyperus L. by Kern (loc. cit.) and
by Koyama (Journ. Fac. Sci. Univ. Tokyo, Sect. 3, 8 : 72 (1961)).

GRAMINEAE
(By W. D. Clayton)
BAMBUSA VULGARIS Schrad. ex Wendl.—Dandy in Exell, Cat.: 365.
ANNOBON : Crater Lake, south side, 265 m., lakeside vegetation, near clearing
in forest, 12 Aug. 1959, Melville 276.
New record for Annobon. Introduced.

PANICUM BREVIFOLIUM L.—Dandy in Exell, Cat.: 370.

ANNOBON : Pico do Fogo, S.W. side, 390 m., bare soil on rocky hillside, 25 July
1959, Melville 187.

New record for Annobon.

OPLISMENUS HIRTELLUS (L.) Beauv.—Dandy in Exell, Cat.: 373.—Exell, Suppl.: 51.
ANNOBON : Path to Quioveo (Pico del Centro), just above north end of Crater
Lake, 300 m., pathside in wood, 18 July 1959, Melville 163.
New record for Annobon.

PASPALUM COMMERSONII Lam.—Dandy in Exell, Cat.: 375.

ANNOBON: N.W. of the Island, 15 m., cassava plantation, 11 July 1959,
Melville 122.

New record for Annobon.

PASPALUM CONJUGATUM Berg.—Dandy in Exell, Cat.: 375.
ANNOBON: Near Ambo, 30 m., streamside, 12 July 1959, Wrigley 28.
New record for Annobon.

ANTHEPHORA CRISTATA (Doell) Hack. ex De Wild. & Dur.—Exell, Suppl.: 52.
ANNOBON: N.E. coast near Ambo, 24 m., pathside in cassava plantation,
21 July 1959, Melville 171.
New record for Annobon,

118 ANGIOSPERMS OF ANNOBON ISLAND

Beckeropsis laxior W. D. Clayton, sp. nov.; affinis B. nubicae (Hochst.) Fig. &
De Not. sed differt ramulis laxioribus subglabris, spiculis acuminatis, glumis
superioribus longioribus, lemmatibus spinuloso-hipsidulis. (Holotype in Herb.
Kew from Nigeria : Summit of Orosun, Idanre Hills, Keay FHI 22678.)
Beckeropsis nubica sensu Dandy in Exell, Cat.: 378. :

ANNOBON : S.W. side of Pico do Fogo, 390 m., bare soil after burning, 25 July

1959, Melville gr.

New record for Annobon. Also in S. Tomé and on the mainland of West Africa
from Ghana to Cameroons.

PENNISETUM POLYSTACHION (L.) Schult——Dandy in Exell, Cat.: 378.—Exell,
Aditam.: 89.
ANNOBON : Ambo, by the expedition’s houses, 15 m., waste ground, common,
13 July 1959, Melville 139.
New record for Annobon.

SCHIZACHYRIUM BREVIFOLIUM (Swartz) Nees ex Buse in Pl. Junghuhn.: 359 (1854).
Andropogon brevifolius Swartz, Nov. Gen. & Sp. Pl.: 26 (1788).
ANNOBON:: N.W. of the Island, 15 m., cassava plantation, 11 July 1959, Melville

128.
New record of the genus Schizachyrium Nees for the islands. S. brevifolium is

widespread in the tropics.

CYMBOPOGON CITRATUS (DC.) Stapf.—Dandy in Exell, Cat.: 380.

ANNOBON : Ambo, just north of the expedition’s houses, fide Melville (no
specimen collected).

New record for Annobon. Introduced.

CoIx LACRYMA-JOBI L.—Dandy in Exell, Cat.: 381.
ANNOBON : Near Ambo, 24 m., streamside, 12 July 1959, Wrigley 21.
New record for Annobon. Introduced.

Bull. B.M. (N.H.) Bot. 3, 3 PLATE 3

Mecklenburg

s 4040
nee GID

Turraea glomerulifilora Harms (Mildbraed 6487, holotype)

Bull. B.M. (N.H.) Bot. 3, 3

"ay Pease aunrlonenagd (Lets %
rita. }

Leelee,
per. CAL E44 19.6 2.

CAMBRIDGE Qa EXPEDITION
TO ANNOBON ISLAND
WEST TROPICAL AFRICA

T. C. Watctey ano F. A. MELVILLE, 1959

ANNORGs Sidee west of Croter Lake,
Open woodland, L200ft. Alte

Flover white; Seaerous frudt ith 1 orange

Maytenus annobonensis (Loes, & Mildbr.) Exell (Wrigley 56, topotype)

” 1, | rex Ey 1 ‘|

PLATE 4

Bull. B.M. (N.H.) Bot. 3, 3

ae | (14 4

PLATE 5

CAMBRIDGE SamBMN EXPEDITION

TO ANNOBON ISLAND
WEST TROPICAL AFRICA
T. (. Wreictey anp FL AL Menyrire, 1959 i
}
J

(igelaca ovat; Sek elbus- :
Der. 2orSacw 1962

AANOBON: Aabo,
Dey val SOOM. Alt,

Tree t 30ft. Fruit red and yellow
With back onl oruye seed, Flowers
off—wiite (:inkieh),

F, ¥elvil 232, 2 Auge 195996

Agelaea ovalis Schellenb. (Melville 232, topotype)

Bull. B.M. (N.H.) Bot. 3, 3 PLA LES

Arb raat

; j i
antl

thee noi Friedrich Herzog zu Mecklenburg

Zweite Expedition nach Central-Afrika 1910-1911 ..

Coots prunes. dene bOntetts OAL de. He

gee pect, ae owes. Aeh ee
a ke

o1 Annobon ; Be é Dtttoad han 2 lon-

byrrnens a. /Fibetarelef Cu PbO a

Cassipourea annobonensis Mildbr. ex Alston (Mildbraed 6511, holotype)

Bull. B.M. (N.H.) Bot. 3, 3

PLATE 7

Tice

CAMBRIDGE @0N@@088) EXPEDITION
TO ANNOBON ISLAND
WEST TROPICAL AFRICA

T. C. Wrratey ann F. A. Menviiie, 1959
ANNOBON: Pieo Surgado,

Forest ULS0ft, Alt.
Flower pale pink,

T.c. Trighley 65, 2k July 1959,

Tristemma oreothamnos Mildbr. (Wrigley 65, topotype)

Bull. B.M. (N.H.) Bot. 3, 3 PLATE 8

WERE ee

ANNGBON: Nr. highest polst of crater.

CAMBRIDGE SRBRGSRY EXPEDITION

TO ANNOBON ISLAND Forest. 1600ft, Alt.
WEST TROPICAL AFRICA Undershrub. 2 speciuens with white corollas;
A others fruit oly.
© Gh . A. Mr Ey 195 igl
T. C. Warcrey anno F. A. Mrivinre, 1959 T.C. te 93. 26 July, 1959.

Bertiera annobonensis G. Taylor (Wrigley 93, topotype)

Bull. B.M. (N.H.) Bot. 3, 3 PLATE 9

Po

Se * rare:
te bntn hilt
2 ay ee te . Lda
ye Souter iets en

POST: ae Pare pte Tr tacth yy

on Dr. J. Mildbraed. No? O% ; oo al

eo:

Nuxia congesta R. Br. ex Fresen. var. congesta (Mildbraed 6561, holotype of Lachnopylis
annobonensis Mildbr.)

Bull. B.M. (N.H.) Bot. 3, 3 PLATE 10

CAMBRIDGE QR EXPEDITION
TO ANNOBON ISLAND
WEST TROPICAL AFRICA

T. C. Wrisctey ann F. A. Mervinne, 1959

Thecacoris annobonae Pax & Hoffm. (Wrigley 92, topotype)

Bull. B.M. (N.H.) Bot. 3, 3 PLATE 11

‘Diseeclaswylorn pubescens Pars 5 , tof,
hee

Der, Oe Seek i Fee
AR BON, hichest poet of crater. CAMBRIDGE S2a¥BRS@8% EXPEDITION
orest. ieoort, Alt. TO ANNOBON ISLAND
Lurye leaves 2/ phyllotetigs, Catkin-like WEST TROPICAL AFRICA
infior-scses, heacreus grees fruits,
7. trisley 96, a6 Taly 1959, T. C. Wrigiey ann F. A. Mrtvitee, 1959

Discoclaoxylon pubescens (Pax & Hoffm.) Exell (Wrigley 96, topotype)

Bull. B.M. (N.H.) Bot. 3, 3

PLATE 12

ie Be a 1

A. OBON: Riige west of Crater Lake,
Open woodland, 1200ft, alt,
Q & ¢ flowers on same shrub. Low shrub
T.o, “rigley 55, 18 July 1959,

CAMBRIDGE SRRSBIR EXPEDITION
TO ANNOBON: ISLAND
WEST TROPICAL AFRICA

T. C. Wriatey ano F. A. Metyinre, 1959

teal ha auvstnat fan vt Mifhin.
per, Qto-fnta 19 6%

Acalypha annobonae Pax & Hofim. (Wrigley 55, topotype)

A REVISION OF THE GENUS _
PETRORHAGIA ( >

& <s

P. W. BALL
AND

V. H. HEYWOOD

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)

BOTANY Vol. 3 No. 4
LONDON: 1964

es

7S
tes
5,

A REVISION OF THE GENUS PETRORHAGIA

BY

PW, BALL. ang. Veo; HEYWOOD

x wet
(University on verpool)

Php. 119-172 ; 22 Text-figures ; Plates 13-15

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 3 No. 4
LONDON : 1964

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), <‘nstituted in 1949, 1s
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
veady. Volumes will contain about three or four
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within one calendar year.

This paper is Vol. 3, No. 4 of the Botany series.

© Trustees of the British Museum (Natural History), 1964

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Issued September, 1964 Price Twenty-two Shillings

ACREVISION -OF THE GENUS PE FRORAAGCIA

By P. W. BALL and V. H. HEYWOOD

THIS revision was undertaken in connexion with the preparatory work for Flora
Europaea. The genus Petrorhagia (Tunica auct.) had not been the subject of a
recent survey and several interesting problems of generic circumscription required
a solution.

The greater part of this study has been based on herbarium material from the
institutions listed below. Several species are still known from only a few collections
and the scope for field work in this genus is very wide.

Over fifty taxa within the genus have been described at specific level (under
Tumica, Gypsophila, Dianthus, etc.). Twenty-five species are recognized in this
revision.

Numerous taxonomic and distributional problems remain to be solved, especially
in the eastern Mediterranean region where complex variation patterns are evident
from the material it has been possible to examine. These patterns bear some
resemblance to the kind described by Ehrendorfer (1958) as Fossil Hybrid Complexes
in his studies on Galium sect. Jubogalium. Much further factual information is
needed, however, before the possible evolutionary relationships of the species of
Petrorhagia can be outlined.

The species and subspecies are delimited in this revision in a morphological-
geographical sense (cf. Heywood, 1958).

We wish to acknowledge with gratitude the kindness of the Directors or Curators
of the following Herbaria for allowing us to borrow or consult their material (the
abbreviations are those used in the citation of specimens).! Our thanks are due also
to Mr. J. E. Dandy for his assistance with several points of nomenclature.

BM = British Museum (Natural History).
CGE = Botany School, University of Cambridge.

E = Royal Botanic Garden, Edinburgh.

FI = Herbarium Universitatis Florentinae, Florence.

K = Royal Botanic Gardens, Kew.

LIVU = Hartley Botanical Laboratories, University of Liverpool.
W = Naturhistorisches Museum, Vienna.

W-HAL = Halacsy Herbarium, Naturhistorisches Museum, Vienna.

W-RECH = Rechinger Herbarium, Naturhistorisches Museum, Vienna.

1Only a selection of material has been cited under Petrorhagia saxifraga, P. prolifera and P. velutina.
y g 8

122 REVISION OF THE GENUS PETRORHAGIA

CHARACTERS OF TAXONOMIC IMPORTANCE
(1) Habit and duration.

All species of Petrorhagia are herbaceous—either annual or perennial, the latter
often more or less caespitose and woody at the base, with non-flowering rosettes.
Annual species occur in Sect. Pseudotunica subsect. Creticae, Sect. Pseudogypsophila,
Sect. Dianthella and Sect. Kohlrauschia, while perennial species make up Sect.
Pseudotunica subsect. Illyricae and Sect. Petrorhagia. The annual habit was one of
the characters employed to separate Kohlrauschia asa genus from Tunica auct.
but as noted elsewhere it is also found in other sections.

The stems range from simple, unbranched, through slightly branched to much
branched.

(2) Leaf venation.

The number of veins in the leaves ranges from one to five. This number is
usually constant for a species and throughout certain sections such as Sect.
Pseudotumica. On the other hand the leaves in Sect. Petrorhagia have from one to
five veins, variation even occurring within a single species.

(3) Inflorescence.

The inflorescence is extremely variable in the genus, although it is often constant
within a section or subsection. The inflorescence is basically a dichasium, but in
some species it is very condensed, so that the flowers are fasciculate, while in Sects.
Dianthella and Kohlrauschia and Sect. Petrorhagia subsect. Thessalae a capitulum
is formed with large outer bracts enclosing the rest of the inflorescence.

In addition to the bracts associated with branches of the inflorescence a few species
always have two or more pairs of “ epicalyx bracts’’ inserted just beneath the
calyx of each flower and more or less enclosing it. These are most easily seen in
P. saxifraga. In species with a fasciculate or capitate inflorescence it is not always
possible to decide whether such epicalyx bracts are present or not. It must be
emphasized that these structures are not epicalycine in nature since they do not
form part of the calyx proper.

The consistency and colour of the bracts vary from membranous to coriaceous
and from colourless to brown or purple, and the shape from lanceolate to suborbicular.

(4) Calyx.

The calyx is 5-toothed, and varies from obconic to cylindric in shape, and is
usually thin and membranous. Each tooth has I or 3 (rarely more) veins associated
with it, forming a costa of the calyx ; there are 5 costae altogether. Between
each pair of costae is a thin membranous, veinless commissure. In Sect. Dianthella
the costae are very broad and often meet laterally so that the commissures are
more or less absent (Figs. 19, 20).

The central vein of each costa is usually quite distinct but, in those species with
3 veins, the laterals are sometimes very weak and indistinct. The number of veins
is sometimes constant within a section (e.g. Sect. Kohlrauschia), but may vary
within a single species (e.g. P. illyrica).

REVISION OF THE GENUS PETRORHAGIA 123

There are two main types of calyx teeth in the genus. The first, found in Sects.
Pseudotumica, Pseudogypsophila and Dianthella and in two species of Sect. Petrorhagia,
has a strong central vein reaching the apex of the tooth and often forming a mucro.
The second, found in Sect. Kohlrauschia and the rest of Sect. Petrorhagia, has all
the veins disappearing beneath the apex of the tooth, the apex usually being obtuse.
P. fasciculata, in Sect. Petrorhagia, is somewhat intermediate between these two
types, with a central vein scarcely reaching the apex of a subacute tooth.

(5) Petals.

The petals of most species are linear- or oblong-oblanceolate with an entire
rounded apex and without a distinct claw. Sect. Kohlrauschia is unusual in that
the petals have a distinct long claw and usually an obcordate limb, although
P. glumacea var. glumacea has a deltate limb with an irregularly toothed apex.
However, the species of Sect. Petrorhagia subsect. Thessalae have more or less
distinctly clawed petals, with a small elliptic-orbicular limb, while some species in
Sect. Petrorhagia subsect. Saxifragae have obcordate petals which are not distinctly
clawed. One or two species in other sections also have retuse or shortly 2-lobed
petals while P. hispidula has 4-lobed petals.

(6) Seeds.

The seed structure is identical with that found in Dianthus and Velezia and it
appears to be unique within the family. The embryo is straight and the seeds are
dorsiventrally compressed usually with incurved or thickened margins, the hilum
being situated in the centre of the concave surface. There is considerable variation
within the genus but two main types can be recognized. One is found only in
Sect. Pseudotunica, which has relatively large seeds (usually c. 2 mm. long or longer)
which have thin margins. They are black when mature and almost smooth. The
remaining sections have usually smaller seeds (0-8-1-5 mm. long) with distinctly
thickened margins. They are dark or blackish-brown when mature, and distinctly
reticulate or tuberculate or even papillose on the convex surface. In Sect. Pseudo-
tunica, P. lycica and P. candica are somewhat intermediate between the two extremes.

(7) Cytology.
The following chromosome counts have been recorded in Petrorhagia:
Sect. Petrorhagia
P. saxifraga gw == Go: «ch Love c: Love (1961)
Sect. Pseudogypsophila
P. alpina subsp. olympica 2n = 30 cf. Darlington & Wylie (1955)
Sect. Kohlrauschia

P. nanteuilit an = 00. .cl, Love & Love (1961):
Ball & Heywood (1962)
P. prolifera an = 30. cl. Love & Love (1961).,
Ball & Heywood (1962)
P. velutina 2n = 30 cf. Darlington & Wylie (1955) ;

Ball & Heywood (1962)

124 REVISION OF THE GENUS PETRORHAGIA

The number for P. velutina has been confirmed in material from Greece, and this
also possesses the short chromosome noticed by Bocher e¢ al. (1953) in material
from Portugal. The difficulty of obtaining seed of most other species has prevented
any further cytological study of the genus. The basic number is x = I5 as in
Dianthus, but different from Gypsophila where according to Barkoudah (1962)
it is xX = 17 or 18. Polyploidy occurs in Sect. Kohlrauschia, cf. Bocher et al. (1953,
1955), Ball & Heywood (1962), and it may also occur in other parts of the genus,
especially in the eastern Mediterranean region where a considerable degree of
hybridization is occurring or has occurred in the past.

(8) Experimental studies.

Studies on the variation and physiological responses of samples of P. prolifera,
P. velutina and P. nanteuilia grown from seed of various provenances have been
made by Bocher e# al. (1953, 1955).

(9) Ecology.

The evidence available from collectors’ notes on herbarium labels, from Floras
and from limited personal observations shows that most species of Petrorhagia
grow in dry, sandy or calcareous habitats where there is little competition from other
species. They may be found in sand, rocky debris, on rocks and cliffs, detritus,
roadsides, walls, dry grassland, etc. P. alpina subsp. olympica frequently occurs
in montane coniferous woodland, sometimes at the sides of streams, so it would
appear that at least this subspecies prefers a relatively moist and shaded situation.

(10) Geographical distribution.

The genus occurs mainly in the eastern Mediterranean region and south-eastern
Europe, extending west to Madeira and the Canaries (one species) and east to
Pakistan and Kashmir (one species). A few species occur in North Africa. With
the exception of P. saxifraga, P. prolifera and P. nanteuili1, the genus is absent
from east, west, central and north Europe.

The greatest concentration of species occurs in Greece and Turkey with 12 and g
species respectively. Eight species or subspecies are endemic to Greece and
neighbouring Aegean islands, two to Crete, four to Turkey, one to Iran, one to
Cyprus and three to North Africa.

There are four widespread species in the genus—P. saxifraga, which occurs
throughout much of central and southern Europe and extends into south-west Asia ;
P. prolifera, which is distributed in central Europe and in the mountains of southern
Europe, the Caucasus, northern Anatolia, and in those of western North Africa ;
P. velutina, which is widespread in the Mediterranean region ; and P. alpina,
which is found in the mountains of central and western Asia, extending to southern
Bulgaria.

RELATIONSHIPS AND DELIMITATION OF PETRORHAGIA WITHIN
THE SILENOIDEAE
A survey of the relationships of Gypsophila and segregate genera and their
distinction from Twnica (i.e. Petrorhagia) is given in Barkoudah’s recent revision
(1962). As noted there, Linnaeus included two species of Tunica in his treatments

REVISION OF THE GENUS PETRORHAGIA 125

of Gypsophila, possibly failing to distinguish between the two groups on account
of the numerical basis of his Sexual System. Haller (Enum. Stirp. Helv. 1 : 381
(1742) ; Hist. Stirp. Helv. 1 : 39 (1768)) “‘ made the boundaries between Gypsophila
and Tunica sufficiently clear’”’, as Barkoudah says, but in fact Haller was using
the name Tunica in the sense of Dianthus.

Linnaeus (Sp. Pl., 1753) included two species of Petrorhagia in Dianthus—
D. saxifragus (=P. saxifraga) and D. prolifer (=P. prolifera), and a further one in
Gypsophila—G. rigida (=P. saxifraga var. glomerata). In 1759 (Syst. Nat., ed. 10)
he transferred D. saxifragus to Gypsophila. Linnaeus (1763 and 1767) and Arduino
(1764) described two further species (P. cretica and P. tllyrica) under Saponarta.
A number of other species were subsequently described under Gypsophila or Dianthus,
but the genus Tunica was not recognized as independent until Mertens & Koch
(1831) and Fischer & Meyer (1837) redefined it, at the same time misapplying
the name. These latter authors included in Tunica all the species of Petrorhagia
known at that time, except for those of Sect. Kohlrauschia which were still retained
in Dianthus. Kunth (1838) subsequently separated Kohlrauschia from Dianthus
as a distinct genus. Boissier in his Flora Orientalis reviewed all the known species
and concluded that Kohlrauschia was best united with Tunica and his view was
followed by Bentham in Benth. & Hook., Gen. Pl. 1 : 145 (1862)), by Pax (in Engler
& Prantl, Nat. Pflanzenfam. 3 (1b) : 76 (1889)) and by Pax & Hoffmann (in
Engler & Prantl, op. cit., ed. 2, 16c : 355 (1934)). Most modern European Floras,
however, still recognize Kohlrauschia and Tunica as distinct genera, but only one
of these, Hayek (Prodr. Fl. Penins. Balcan. I : 221-224 (1924)), treats of more than
three or four species. This is an interesting example of the role of historical and
local influences affecting taxonomic treatment.

As circumscribed in this revision Petrorhagia is more or less intermediate in most
of the taxonomically significant characters between Dianthus and Gypsophila.
It resembles Gypsophila in the following features :

(1) the calyx nearly always with membranous veinless commissures between
the teeth ;

2) the calyx usually 5- or 15-veined ;
) the flowers sometimes ebracteate ;

) the petals usually not clawed ;

) in inflorescence and general appearance (Sects. Pseudotunica and Pseudo-
gypsophila).

In the following characters it resembles Dianthus :

(
(3
(4
(5

)

) flowers sometimes bracteate ;

) petals sometimes clawed and with a broad limb (Sect. Kohlrauschia) ;

) in inflorescence and general appearance (Sect. Kohlrauschia) ;

) seed structure—seed compressed dorsiventrally with the embryo straight,
lying lengthwise in the middle, not compressed laterally with the embryo
curved and peripheral.

126 REVISION OF THE GENUS: PETRORAAGIA

There is no doubt, however, of the much closer relationship on general grounds of
Petrorhagia to Dianthus than to Gypsophila. Of the characters listed above only
one, the seed structure, is constant and reliable. In all the features in which
Petrorhagia resembles Gypsophila there is an intergradation with the opposing
Dianthus character. Thus in characters (1) and (2) the membranous commissures
are virtually absent in P. pamphylica while the same species has 5 or 7 veins to
each costa. This species is linked to the rest of the genus through P. peroninit.
In character (4) Sect. Kohlrauschia has a distinct long claw with a broad limb,
Sect. Petrorhagia subsect. Thessalae has a long, more or less distinct claw but a small
limb, while in Sect. Petrorhagia subsect. Saxifragae some species (e.g. P. graminea)
have petals with a broad limb but virtually noclaw. There is thus almost a complete
range from the Gypsophila type of petal found in Sects. Pseudotunica, Pseudo-
gvpsophila, Dianthella and part of Sect. Petrorhagia to the Dianthus type found in
Sect. Kohlrauschia.

Characters (3) and (5) are more complex. Gypsophila normally has a large,
much-branched panicle with numerous small flowers, sometimes aggregated into
small heads or fascicles. All the bracts are usually small and are only associated
with branches of the inflorescence. Dianthus, on the other hand, usually has a
little-branched inflorescence with the flowers solitary, fasciculate or capitate at the
tips of the stems. It frequently has large brown scarious bracts surrounding
solitary flowers or fascicles or capitula. Both situations are found in Petrorhagia.
Sects. Pseudotunica and Pseudogypsophila and Sect. Petrorhagia subsect. Saxifragae
closely resemble Gypsophila. However, P. saxifraga in Sect. Petrorhagia subsect.
Saxifragae has bracts at the base of solitary flowers, and which are clearly not
associated with branches of the inflorescence. The Dianthus situation is found in
Sect. Kohlrauschia which is almost unbranched with terminal heads of flowers
surrounded by large brown scarious bracts. The remaining two groups, Sect.
Dianthella and Sect. Petrorhagia subsect. Thessalae, are more or less intermediate
between the two extremes. Sect. Dianthella has numerous brown or purplish
scarious bracts surrounding one or a few flowers, although the stems are normally
much branched. Sect. Petrorhagia subsect. Thessalae has little-branched stems
with small terminal capitula. The bracts are usually relatively small and mem-
branous except for a stout mid-vein, but in one species, P. thessala, they tend to
be larger and broader with a broad brown scarious region.

It will be seen from this survey that a clear assessment of the affinities of the
genus can only be obtained from the complex of characters referred to as the seed
structure. Here there is no gradation with the Gypsophila type. Seed structure,
in fact, distinguishes Petrorhagia, Dianthus and Velezia from all other genera of the
subfamily Szlenoideae. This fact lends strong support to the grouping of these
three genera together into a distinct subtribe as is done by Bentham (in Benth. &
Hook., Gen. Pl. 1 : 142 (1862)) rather than to the arrangement of Pax (1889) and
Pax & Hoffmann (1934), in which Tunica is placed with Gypsophila and Dianthus,
and Velezia with Saponaria and Vaccaria.

The distinction between Petrorhagia (Tunica) and Velezia was discussed fully

REVISION OF THE GENUS PETRORHAGIA 127

by Davis (1957). The inclusion of Kohlrauschia within Petrorhagia does not sub-
stantially alter any of Davis’s arguments or conclusions.

The characters by which Kohlrauschia, when treated as a separate genus, is
usually separated from Petrorhagia (Tunica) are as follows :

(1) annual ;
(2) inflorescence a capitulum ;
(3) petals with a long claw and a distinct, broad limb.

Annuals are found in three other sections of Petrorhagia while the unreliability of
the remaining two characters has been discussed above. If Sect. Kohlrauschia is
treated as a distinct genus, then there can be little justification for not treating
Sects. Dianthella and Pseudotunica as distinct genera as well.

The expressions of the more important sectional characters throughout the genus
are shown in the accompanying table. The sections are relatively distinct although
the position of two species, P. kennedyae and P. phthiotica, is still not clear. This
is discussed more fully under the species in question.

REVISION OF THE GENUS PETRORHAGIA

128

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jo 1oquinyny

REVISION OF THE GENUS PETRORHAGIA 129

PETRORHAGIA (Ser.) Link

PETRORHAGIA (Ser.) Link, Handb. Erkenn. Gewiachse 2 : 235 (1831).

Imperatia Moench, Meth. Pl. : 60 (1794) ; non Imperata Cyr. (1792).

Gypsophila sect. Petrorhagia Ser. in DC., Prodr. 1 : 354 (1824).

Tunica sensu Mert. & Koch, Rohl. Deutsch. Fl. 3 : 182 (1831).—Fisch. & Meyer, Index
Sem. Hort. Bot. Petrop. 4: 48 (1837).—A. Braun in Flora 26 : 384 (1843).—Boiss., FI.
Or. I : 516 (1867).—Pax in Engler & Prantl, Nat. Pflanzenfam. 3 (1b) : 76 (1889).—
Pax & Hoffm. in Engler & Prantl, op. cit., ed. 2, 16c : 355 (1934), et auct. plur.; non
Tunica Ludw. (1757).

Gypsophila subgen. Tunica Reichb., Fl. Germ. Excurs. : 802 (1832).

Kohlrauschia Kunth, Fl. Berol. Fam. Nat. Dispos. 1 : 108 (1838).

Dianthus sect. Kohlrauschia (Kunth) Fenzl in Endl., Gen. Pl. : 971 (1840).

Dianthus sect. Tunica (Reichb.) Fenzl, loc. cit. (1840).

Fiedleria Reichb., Deutsche Bot. 1 : 206 (1841) ; Ic. Fl. Germ. & Helv. 6: 42 (1844).

Tunica subgen. Imperatia Jaub. & Spach, Ill. Pl. Or. 1: 11 (1842).

Dianthella Clauson ex Pomel, Mat. Fl. Atlant. : 9 (1860).

Tunica sect. Fiedleria (Reichb.) Graebn. in Aschers. & Graebn., Synops. Mitteleur. F1.

5 (2) 3. 292. (2921).

Type) Pusaxifraga (L:) Link:

Annual or perennial herbs, sometimes with a woody stock. Leaves opposite,
subulate to oblong, I-3(-5)-veined, entire or the margin minutely serrulate.
Inflorescence a dichasial cyme, with the flowers solitary, fasciculate, or capitate ;
bracteate or ebracteate, sometimes with decussate bracts surrounding the calyx
(epicalyx bracts). Calyx obconic to cylindric, 5-toothed, each tooth with I or 3
(rarely up to 7) veins and usually with broad membranous veinless commissures
between the teeth. Petals 5, clawed or not, the apex entire to bifid, rarely 4-lobed
or irregularly toothed ; without coronal scales. Stamens 10. Styles 2. Capsule
opening by 4 apical teeth. Seeds numerous, dorsiventrally compressed, scutate ;
embryo excentric, straight.

The genus Petrorhagia as interpreted in this revision includes the species previously
included by most authors in the genera Tunica and Kohlrauschia. A wide circum-
scription, similar to the one adopted here, was first used by Boissier, Fl. Or. 1 :
516-523 (1867).

The name Tunica has been attributed variously to Haller, Scopoli and Koch.
It was in fact first validly published by Ludwig, Inst. Reg. Veg., ed. 2 : 129 (1757),
as an illegitimate substitute for Dianthus L. (see Dandy, 1957). Haller (1768) and
Scopoli (1772) also used Tunica as a replacement for Dianthus, all the species recog-
nized by these authors being originally described by Linnaeus under Dianthus.

Mertens & Koch (1831) appear to have been the first authors to treat Tunica as
a genus distinct from Dianthus. They recognized only one species, 7. saxifraga
(L.) Scop. Subsequent authors, e.g. Boissier (1867) and Williams (1890), followed
Mertens & Koch’s misuse of the name Tunica, although Williams himself quite
clearly states that Tunica of Scopoli is a replacement for Dianthus. Janchen (1963)
likewise proposes acceptance of Tunica of Scopoli as emended by Koch, but even
if this were possible the name would still be a later homonym of Tunica Ludwig,
itself a synonym of Dianthus. Maire (1963) goes further and cites “ Tunica Boehm.

130 REVISION OF THE GENUS PETRORHAGIA

in Ludw. (1760),nom. abort., emend. Mert. et Koch (1831)’’ as a nomen conservandum,
but as far as we are aware no such proposal has been made.

The typification of Petrorhagia requires some consideration. Link placed under
it four species, P. glomerata, P. multicaulis, P. rigida and P. saxifraga. P. glomerata
is currently treated as a Gypsophila, the remaining species being referred to Tunica
or Kohlrauschia. It is therefore necessary to typify the name Petrorhagia since
Link himself indicated no type. Link’s genus is based, by implication, on Gypsophila
sect. Petrorhagia Ser. in DC., Prodr. 1 : 354 (1824). He did not directly cite
Seringe or De Candolle, but his genus contains the same species as Seringe’s section,
except for G. dianthoides, which Seringe cited with a question mark. Accordingly
we have to typify Gypsophila sect. Petrorhagia Ser. The definition given by Seringe
s ““ Calycis squamis scariosis 2—4 oppositis basi instructi’’. This agrees well with
G. saxifraga which normally has 2 opposite pairs of bracts inserted just below the
calyx and more or less completely enclosing it. Seringe again refers to these bracts
in the description of this species, and a further point, which clearly indicates that
Seringe regarded it as typical, is that “‘ petrorhagia’”’ is a simple translation into
Greek of “saxifraga’’. The typification of Petrorhagia by P. saxifraga was pro-
posed ‘by Britton & Brown (Ill. FIN. U.Ss& Can. ed. 2; 22:72: (19%3)) and there
seems no good reason to reject it.

Dandy (1957), unaware of the previous typification by Britton & Brown, proposed
Gypsophila glomerata as the type of Sect. Petrorhagia Ser. and consequently of Link’s
generic name. In G. glomerata there is sometimes a pair of opposite bracts towards
the base of the pedicel of the lateral flowers of a partial dichasium, but they are
always clearly separated from the base of the calyx by a length of pedicel, and the
central flowers never have such bracts. This interpretation of the structure of
G. glomerata agrees with the inflorescence diagram given by Barkoudah (1962) in
fig. 1 of his revision of Gypsophila. In our view the bract situation in G. saxifraga
agrees better with Seringe’s definition than does that of G. glomerata. Barkoudah,
apparently following Dandy, gives Petrorhagia as a synonym of Gypsophila. This
according to our view is not correct, except in so far as P. glomerata is concerned.

KEY TO THE SPECIES

Bracts enclosing the calyx ; inflorescence often capitate-fasciculate :
Petals with a long distinct claw and a broad obcordate or obdeltate limb, pink or
purplish ; largest bracts usually at least 4 mm. broad :
Seeds not more than 1:3 mm. long, papillose ; leaf sheaths at least twice as
long as broad . 2 é 24. P. velutina
Seeds at least 1-3 mm. long, alan to tuberculate : leaf sheaths not more
than twice as long as broad :
Petal-limb crenate or laciniate ; seeds 1-7—2:2 mm. long, almost smooth or
sparsely tuberculate ; : : . 25. P. glumacea
Petal-liimb obcordate ; seeds 1-3-1-8 mm. long, reticulate or tuberculate :
Seeds reticulate : : A ; ; ; . 22. P. prolifera

REVISION OF THE GENUS PETRORHAGIA 131

Seeds tuberculate :
Outermost bracts of the head mucronate (western Europe and western
North Africa) : ; . 23. P. nanteuilit
Outermost bracts obtuse (Balkan peninsula) , . 25. P. glumacea
Petals not distinctly clawed or if clawed then with a small elliptic limb, white or
pink ; largest bracts usually less than 4 mm. broad :
Leaves 3-veined ; annual, rarely perennial :
Bracts I-veined ; perennial . ; ; : ; . 12. P. fasciculata
Bracts 3- to many-veined ; annual :
Costae of calyx 5- to 7-veined ; seeds 1-6-2 mm. long, 1:2-1°7 mm. broad
, 20. P. pamphylica
Costae of calyx 3-veined ; seeds I-I-1-2 mm. long, 0-7—0-9 mm. broad
21. P. peroninit
Leaves I-veined or obscurely veined ; perennial :
Flowers solitary or fasciculate ; bracts and epicalyx bracts distinctly shorter
than the calyx ; stems usually much branched :
Stems glandular-pubescent at least at the base ; calyx teeth more or less

triangular, acute or subobtuse ; . 12,.P. fasciculata

Stems eglandular ; calyx teeth oblong, obtuse : i, (DBE SARL PAR
Flowers capitate ; bracts more or less completely enclosing the calyx :

Calyx teeth more or less triangular, acute (Crete) 19. P. dianthoides

Calyx teeth oblong, obtuse :
Petals white with pink or purple veins ; outer bracts brown-scarious
with white-membranous margin : : ‘ 16. P. thessala
Petals white without coloured veins ; outer bracts almost entirely

white-membranous :
Stems glabrous or papillose-scabrid at the base ; largest bracts
I-5—2:5 mm. broad ; es 5°5-8 mm. long, pale-red on the lower

surface d ‘ 18. P. macra
Stems pubescent at the base: largest bracts 0: 7 1:3 mm. broad ; petals
8-9 mm. long, not red on the lower surface . 17. P. cyrenatca

No bracts enclosing the calyx ; flowers solitary, rarely a few fasciculate :
Petals emarginate, retuse or 4-lobed at the apex :

Petals 4-lobed F ; 5 . : P : . 8. P. hspidula
Petals emarginate or retuse :
Calyx teeth oblong, obtuse ; petals usually pink or reddish, or white or
yellow with pink or purple veins :
Basal leaves 8-20 mm. long ; petals pie mm. broad, obcordate, pink,

reddish or white : . 14. P graminea
Basal leaves 4-8 mm. long ; peta o-8-I mm. ‘broad, pale yellow with
purple veins : . 15. P. rhiphaea

Calyx teeth triangular-oblong, more or - Jess ‘acute or mucronate ; petals
white, sometimes with pink veins :
Costae of calyx 3-veined (the lateral veins sometimes weak near the apex) ;

132 REVISION OF THE GENUS PETRORHAGIA

seeds c. 1-5 mm. long, black, smooth . , : 4. P. candica
Costae of calyx I-veined ; seeds c. I mm. long, blackish-brown, more or less
tuberculate ; ; ; : : : . 12. P. fasciculata

Petals entire”
Annual, without non-flowering rosettes :
Costae of calyx 3-veined ; seeds 2-3-5 mm. long, smooth, with thin margin :
Calyx glabrous ; petals included in the calyx or rarely shortly exserted ;

seeds 2-2-8 mm. long . ? ; Or, cretion
Calyx densely glandular-pubescent ; petals distinctly exserted from the
calyx ; seeds 2:7-3-5 mm. long . ; ; 7. P. arabica

Costae of calyx I-veined ; seeds less than 2 mm. one, reticulate-tuberculate,
with thickened margin :
Stem glabrous ; petals 3-5-6 mm. long, entire. g. P. alpina
Stem viscid-glandular at the apex ; petals 6-8-5 mm. ieee bifid
10. P. kennedyae
Perennial, more or less caespitose and woody at the base, with non-flowering
rosettes :
Costae of calyx 3-veined :
Lower leaves 15-30 mm. long ; petals often purple-spotted at the base ;

seeds 1-8-2: mm. long, I-1-3 mm. broad __. 3. P. armerioides
Lower leaves 10-15 mm. long ; petals purple-veined on the lower surface ;
seeds 1-5 mm. long, 0-5 mm. broad : , : ~~ Sul. bvCICe

Costae of calyx I-veined :
Petals pink ; seeds blackish-brown, reticulate-tuberculate
11. P. phthiotica
Petals white or yellow ; seeds black, smooth :
Stem densely glandular-pubescent throughout, or glabrous in the lower
part, rarely completely glabrous ; ace white or pale yellow ;

anthers usually purple. ; : 1 a ulyrica
Stem densely glandular-pubescent at ‘he base, usually glabrous in the
upper part ; petals yellow ; anthers white 2. P. ochroleuca

Sect. 1. PSEUDOTUNICA (Fenzl) Ball & Heywood

Petrorhagia sect. Pseudotunica (Fenzl) Ball & Heywood, comb. nov.

Dianthus sect. Pseudotunica Fenzl in Endl., Gen. Pl. : 971 (1840).
Tunica subgen. Pachypleura Jaub. & Spach, Ill. Pl. Or. 1: 11 (1842).
Tunica sect. Pseudosaponaria A. Braun in Flora 26 : 384 (1843), nom. tllegit.
Gypsophila sect. Tunicastrum Griseb., Spicil. Fl. Rumel. 1 : 184 (1843).
Tunica sect. Gypsophiloides Boiss., Fl. Or. 1 : 519 (1867) ; non T. sect. Gypsophiloides
Griseb. (1843).
Tunica sect. Pachypleura (Jaub. & Spach) Boiss., tom. cit. : 522 (1867) (err. “ pacypleura ’’).
Tunica sect. Pleurotunica F. N. Williams in Journ. of Bot. 28 : 198 (1890), nom. illegit.
Type (lectotype) : Tunica illyrica (Ard.) Fisch..& Meyer (=P. illyrica (Ard.)
Ball & Heywood).

REVISION OF THE GENUS PETRORHAGIA 133

Annual or perennial. Leaves 3-veined. Epicalyx bracts absent. Petals not
clawed. Seeds large, black, smooth, with thin margin.

Subsect. a. Illyricae Ball & Heywood

Petrorhagia subsect. Illyricae Ball & Heywood, subsect. nov.; subsectio typica
sectionis Pseudotunicae, continens species perennes, seminibus 1:5—2-3 mm. longis.
Type: P. illyrica (Ard.) Ball & Heywood.

Perennial. Seeds 1-5—2:3 mm. long.

1. Petrorhagia illyrica (Ard.) Ball & Heywood, comb. nov.
Saponaria illyrica Ard., Animadv. Bot. Specim. Alt. : t. 9 (1764).—L., Syst. Nat., ed. 12,
2: 267-6t Mant. Pl «<70:(5767).
Gypsophila illyrica (Ard.) Smith, Fl. Graec. Prodr. 1 : 281 (1809).
Tunica illyrica (Ard.) Fisch. & Meyer, Index Sem. Hort. Bot. Petrop. 4: 49 (1837).—
Boiss., Fl. Or. 1 : 520 (1767).—Hayek, Prodr. Fl. Penins. Balcan. 1 : 222 (1924).
Fiedleria illyrica (Ard.) Reichb., Ic. Fl. Germ. & Helv. 6 : 42 (1844).

Stem up to 40 cm. high, much branched, densely glandular-pubescent to glabrous.
Lower leaves 10-30 mm. long, linear-subulate to linear-oblong, 3-veined. In-
florescence lax or subfastigiate. Calyx 3:5-6(-6:5) mm. long, glabrous to densely
glandular-pubescent ; costae I- or sub-3-veined ; teeth obtuse and mucronate to
acuminate. Petals 5-5-10 mm. long, oblong-spathulate, entire, white, rarely pale-
yellow, spotted with purple at the base, sometimes with pink veins on the back.
Anthers usually purple. Seeds 1-7—2:3 mm. long, 0-9-1:3 mm. broad, margin not
thickened.

Distribution : Balkan peninsula, southern Italy (Calabria), Sicily, western
North Africa.

1a. Petrorhagia illyrica subsp. illyrica. (Fig. 1.)

Stem more or less densely glandular-pubescent throughout. Calyx 3°5-5:5 mm.
long, densely glandular-pubescent ; costae I-veined. Petals 6-8 mm. long.

GREECE : Peloponnisos: Kyllene prope Gura, 23 June 1893, Halacsy (W-HAL).
Kyllene prope Trikala, Aug. 1871, Heldreich (BM). Kalawrita, 25 June 1889,
Dorfler 404 (W-HAL) ; 10 June 1926, Bornmiiller 184 (BM ; K). Sterea Ellas :
Parnes, June 1847, Heldreich (E) ; June 1931, Guiol 763A (BM). Parnassus,
July 1855, Heldreich, Fl. Graec. 2984 (K) ; 17 Aug. 1856, Heldreich, Herb. Graec. 95
(BM; K). Oeta, 8 July 1937, Balls & Gourlay 3374 (BM; K). Othrys inter
Lamia et Phourka, 20-21 July 1932, Rechinger fil. 2796 (W-RECH). Amphissa,
May 1862, Mill (K). Timphristos, 22-24 July 1932, Rechinger fil. 2810 (W-RECH).
Near Karpenissi, Timphristos, 16 July 1938, Grebenchitkoff (K). Ypsili Koryphi
prope Vato, 11 July 1906, Maire & Petitmengin 176 (W-HAL).

CRETE : Asprovunna (Levka Ori), 21-22 July 1893, Baldacci 107 (BM ; W-HAL).
Lasithi, 1899, Baldacci 293 (BM) (? locality correct).

The sheet of Safonaria illyrica in the Linnaean Herbarium has two distinct

specimens both of which are somewhat anomalous. One is labelled “ Ard.’”’ and
since it agrees with Arduino’s description and plate it may be regarded as at least

134 REVISION OF THE GENUS PETRORHAGIA

an isotype. It has extremely long pedicels (10-15 mm.), small calyx (c. 4 mm.)
with I-nerved costae, and is densely pubescent throughout. It most closely re-
sembles plants from Sterea Ellas and neighbouring regions of Greece, although the
pedicels are exceptionally long.

For discussion of the specimens from Crete see p. 142.

3 4 ‘

Fics. 1-4. 1, Petrorhagia illyrica (Ard.) Ball & Heywood subsp. illyrica. 2, P. illyrica
subsp. haynaldiana (Janka) Ball & Heywood. 3, P. illyrica subsp. tavgetea (Boiss.) Ball
& Heywood. 4, P. ochroleuca (Smith) Ball & Heywood. a, calyx ; 8, petal. (All x5.)

1b. Petrorhagia illyrica subsp. haynaldiana (Janka) Ball & Heywood, comb.
nov. (ig. 2° Plate 73 a.)

Gypsophila haynaldiana Janka in Osterr. Bot. Zeitschr. 20 : 316 (1870).

Tunica haynaldiana (Janka) Nyman, Consp. Fl. Eur., Suppl. 2 : 57 (1889).

Tunica rhodopea Velen. in Sitzungsber. K. Béhm. Ges. Wiss., Math.-nat. Cl. 1894 (29) : 4
(1895).

Tunica illyrica var. haynaldiana (Janka) Hayek, Prodr. Fl. Penins. Balcan. 1 : 222 (1924).

Tunica illyrica subsp. haynaldiana (Janka) Prodan in Savulescu, Fl. Roman. P.R. 2: 215
(1953).

Tunica illyrica auct. ital. ; non Fisch. & Meyer.

Tunica cretica auct. ital. ; non Fisch. & Meyer.

Tunica ochroleuca auct. ital. ; non Fisch. & Meyer.

Tunica angustifolia auct. ital. ; non Briq.

Tunica compressa auct. ital. ; non Fisch. & Meyer.

REVISION OF THE GENUS PETRORHAGIA 135

Stem glabrous in the lower part, or sometimes completely glabrous. Calyx
(4°5-)5--6 mm. long, usually glabrous ; costae I-veined. Petals 7-10 mm. long.

GREECE : Sterea Ellas ; Elikon (Helicon), Guzol (BM) ; May 1934, Atchley 2007
(K). Thessalia: Oxya supra Kastania, 19-20 July 1893, Haldcsy (W-HAL).
Domokos, 14 July 1956, Rechinger fil. 18129 (W). Malakasi, 13 July 1896, Sintenis
939 (E; K; W-HAL). Pindus prope Chaliki, Heldreich (E). Meteora, supra
Kalabaka, 15-16 July 1884, Heldreich (W-HAL). Mt. Pelio (Volos), 30 July 1882,
Heldreich (W-HAL). Ipiros: Konitsa, c. 1930, Guiol (BM). Dodona (district
Janina), c. 1930, Guzol (BM). Dodona, Olycika, 18 June 1895, Baldacci r4 (BM ;
K ;W-HAL). Peristeri, prope Kalawrita, 14 July 1893, Haldcsy(W-HAL). Pindus,
supra Metsovon, 29 July 1956, Rechinger fil. 18362 (W). Mitsikeli, June-July 1920,
Guiol 763 (BM). Cepelovo (district Zagorion), 1896, Baldacci 291 (BM). Janina,
M. Panajoti Xenos, 1895, Baldacci (BM). Makedhonia : Thessaloniki, July 1857,
Orphanides 925 (E ; W-HAL) ; 11 July 1918, Ramsbottom X27 (BM). Edessa,
28 July—3 Aug. 1932, Rechinger fil. 3241 (W-RECH). Near Karamudli, 29 June
1917, Turrill 445 (BM ; K). Near Akbunar, 27 Aug. 1917, Wilson 104 (BM). Mt.
Elias above Kireckoi, 3 July 1918, Rycroft (BM). Lembet, June 1917, Turner 153
(K). Trigonin (Ostima), near Antartikon, 29 June 1932, Alston & Sandwith 1062
(BM ; K).

ALBANIA : Kudesi (district Valona), 27 June 1892, Baldacci 52 (K ; W-HAL).
Lunxheries Mts., above Cajup, 7 Aug. 1935, Alston & Sandwith 2360 (BM; K).
West of Lake Ohrida, 13 Mar. 1938, Hepburn 83 (K).

JUGOSLAVIA : Makedonija : Mont. Cafa Pan, supre Lin (near Ohridsko Jez),
12 July 1921, Soska (W-RECH). Gjevgjeli (Gevgelija), June 1909, Dimonie (E) ;
June 1927, Rechinger fil. 1535 (W-RECH). Prope Veles, 6 July 1905, Adamovié 180
(W-HAL) ; June 1903, Adamovié (K). Orlovo Brdo prope Krivolak, 18 July 1939, anon.
(K). Treska, June 1905, Adamovic (K). Skopje, 23 June 1937, Thompson (K).
Srbija : Prope Vranja, 15 July 1887, Bornmiiller (W-HAL) ; June 1895, June 1897,
Adamovié (K). Nisch, July 1885, Petrovié (W-HAL) ; June 1885, Magnier rro4
(W-HAL). Mont. Krstilovica, 25 June 1896, Adamovic (BM; E). Pirot, July
1886, Petrovic (W-HAL). Prés de Pordt, Magnier rrogb (W-HAL). Mt.
Pljachavica, 26 June 1896, Adamovic (K). Crna Gora: Mt. Jerinja prope
Andrijevica, Rohlena (K).

ButGariA : Plovdiv: Plovdiv (Philippopolis), July 1871, Janka (W-HAL).
Mt. Bonarschkik, June 1890, Pichler in F. W. Schultz 2812 (BM ; K). Dschendem-
tepe, 20 July 1930, Rechinger fil. 1167 (W-RECH). Gunardzik, 17 July 1897,
Stribrny (E). Kalofer, Sept. 1874, Pichler 19rt (W-HAL). Stanimaka (Asenov-
grad), Gheorghieff 262 (W-HAL) ; June 1904, Bierbach (BM) ; July 1906, Stribrny
' (BM). Papasli (Purvomai), July 1915, Stribyny (CGE ; W-RECH). Khaskovo :
Haskovo (Khaskovo), July 1906, Adamovic (K). Kartal tepe, July 1901, Stribrny
(BM). Sofiya: Dragoman, mont. Golem, 1930, Rechinger fil. 1879 (W-RECH).
North of Dragoman, 2 Aug. 1922, Turrill 674 (K). Mt. Vitosa, prope Marcaevo,
Ir July 1953, Vithodzoevsky r40 (K ; LIVU; W). Kurilo, May 1920, Sétribrny
(CGE ; K). Vyvratsa : Berkowiza, 1897, Uromoff 67 (W-HAL). Dermendere, June

136 REVISION OF THE GENUS PETRORHAGIA

1890, Pichler (K) ; July 1894, Stribyny (BM; E; K). Eli Dere, June 1908,
Stribrny (BM).

RoMANIA : Verciorova, Portam ferream, Degen, Fl. Austro-Hung. 2863 (BM ;
K ; W-HAL ; W-RECH). Inter pagos Verciorova et Gurovoie, 23 July 1887,
10 July 1887, 10 July 1895, Degen 19 (BM ; E ; K ; W-HAL) ; 1887, F. W. Schultz
2420 (BM ; K). Infra Orsova, 28 June 1870, Janka (K ; W-HAL ; isotypes).

ItaLy : Calabria : Circa Reggio, 30 June 1877, Huter, Porta & Rigo 256b (BM ;
CGE ; FI; K; W-HAL); 30 June 1856, Huet du\Pavillon 289 (BM ; CGE ;
FI ; K). Fiumara prope S. Luca, 28 May 1877, Huter, Porta & Rigo 256 (BM ;
CGE ; FI; K) ; June 1808, Rigo in Dérfler, Herb. Norm. 3805 (BM ; E; LIVU).
Pellaro, 25 May 1898, Rigo 344 (W-HAL). Collina di Armo (Reggio), Lacaita (BM).
Stilo, 21 June 1877, Arcangelt (FI). Prope Bova, 8 June 1877, Biondi (FI).
Valanidi, 1850, Pasquale 18 (FI). Monasterace, 16 June 1877, Arcangeli (F1).

SicIEY : Palermo; Ledaro: 10537 (BM; -‘CGE:; FI>, KG: W-HAL) = July 21808;
Ross 112 (BM; E; FI; K). Piano dei Favari, 30 June 1855, Huet du Pavillon
(BM ; CGE; FI; K). Sorrentino, 1854, anon. (FI). Gragia, 1871, Lojacano (FI).
Aetne, 1840, Tornabene (FI). Madonie, Todaro (EK). Monte S. Salvatore : Pizzo di
Palermo et Pizzo Antenna, 15 and 22 July 1873, Strobl (K). Caltagirone, 4 June
1843, Herb. Bunbury (CGE). Prope Troina, 2 July 1855, Huet du Pavillon (BM ;
CGE ; FI; K). Monte Scalone supra Polizzi, 22 July 1874, Strobl (K). Siracusa,
1841, Cassia (FI); Nov. 1929, Corrie & Chase H2287 (K). _Presso Augusto
(Siracusa), Oct. 1911, Vaccari (FI). Caltanissetta, 14 June 1899, July rgor,
Giovanni 308 (FI).

This subspecies is difficult to separate from subsp. zlyrica. The populations in
northern Greece and Macedonia cannot be satisfactorily placed under either
subspecies, although here they are included in subsp. haynaldiana on the basis of the
larger flowers and the stem often glabrous at the base. The populations in Italy
and Sicily are similar and are also included here on the generally more robust habit,
larger flowers, longer pedicels and the lower part of the stem sometimes glabrous.
Tunica cretica, T. ochroleuca, T. compressa and T. armerioides, etc. of Italian authors
are only minor variants of this subspecies.

1c. Petrorhagia illyrica subsp. angustifolia (Poir.) Ball & Heywood, comb. nov.

Silene angustifolia Poir., Voy. Barb. 2 : 164 (1789).

Gypsophila compressa Desf., Fl. Atlant. 1 : 343 (1798).

Tunica compressa (Desf.) Fisch. & Meyer, Index Sem. Hort. Bot. Petrop. 4 : 50 (1837).

Dianthella compressa (Desf.) Pomel, Mat. Fl. Atlant. : 9 (1860).

Tunica davaeana Coss. in Bull. Soc. Bot. France 36 : 103 (1889).—Maire, Fl. Afr. Nord to :
284 (1963).

Tunica angustifolia (Poir.) Briq., Prodr. Fl. Corse 1 : 544 (1910).

Tunica scoparia Pampan. in Arch. Bot. Forli 12 : 25 (1936).—Maire, tom. cit. : 285 (1963).

Tunica illyrica subsp. angustifolia (Poir.) Maire in Bull. Soc. Hist. Nat. Afr. Nord 30 : 265
(1939) ; Fl. Afr. Nord 10 : 283 (1963).

Indumentum variable, stems usually densely pubescent throughout or glabrous
in the lower part. Pedicels strongly compressed (usually more or less terete in all

REVISION OF THE GENUS PETRORHAGIA 137

other subspecies). Calyx 5:5-8 mm. long ; costae I—veined, rarely 3-veined.
Petals 8-11 mm. long.

Morocco : Settat ad Guicer, 31 May 1913, Pitard 2699 (K). Settat, Bir Chafoi,
25 May 1913, Pitard 2697 (BM). Atlas med., Ras-el-Ma prope Azrou, 24 May 1926,
Lindberg 4329 (W). Azrou, 30 June 1923, Jahandiez 628 (BM) ; 23 June 1926,
Lindberg 7072 (K). High Atlas, Tadderte, 28 July 1955, Newbould 48 (BM).
Anamer, Oed Arboa, 16 Aug. 1955, Newbould 136 (BM). Marrakesh-Ouarzazet road,
4 May 1937, Masters 46 (BM). Djebel Afougueur, 9 June 1875, Soc. Dauphinoise
1878, 1113b (K). Province Shedma, May 1871, J. Ball (BM). Ida Oubahil, 1875,
Cosson (K). Shedma, Apr.-May 1871, Hooker (K). Siksona, May 1871, Hooker
(K). Oujda, 21 May 1930, Faure (K). Sidi bou Ziane, 12 May 1913, Pitard 2698
(K).

SPANISH Morocco : Atlas Rifain, valles du Ketama, 21 June 1934, Sennen 9293
(BM).

ALGERIA : Constantine, 18 Aug. 1855, Choulette 22 (CGE; K). Mantourah,
June 1869, E. G. Paris 308 (K). Oran, May 1889, Luzzet (BM). Mostaganem, 15 June
1853, Balansa 138 (K). Ain-Tindamine (Oran), 7 June 1921, Faure (BM). Plaine
de Batna, June 1853, Balansa (K). Crezel vers le Nador de Ciuret, 27 June 1937,
Faure (E). Ain-el-Ibil, Losobria (K).

Tunisia : Djebel Zaghouan, 9 July 1854, Kralik (K).

LisyA : Garian, 16 Nov. 1948, Johnson 54 (BM); 18 May 1952, Guichard
KG/LIB/403 (BM) ; 24 May 1939, Sandwith 2758 (K). Suani Bin Adem, 10 Nov.
1952, Guichard KG/LIB/5 (BM). Benghazi, Grotta di Lete, 1 Apr. 1939, Sandwith
e227 (K).

A variable subspecies with a range of variation parallel to that found in subspecies
allyrica and haynaldiana and in P. armerioides. It can be separated only with some
difficulty from subsp. haynaldiana as the reliability of the pedicel character is
uncertain.

1d. Petrorhagia illyrica subsp. taygetea (Boiss.) Ball & Heywood, stat. nov.
(Fig. 3.)
Tunica illyrica var. taygetea Boiss., Fl. Or. 1 : 521 (1867).
Tunica taygetea (Boiss.) P. H. Davis in Not. R. Bot. Gard. Edin. 22: 165 (1957), excl. syn.
T. cretica et Gypsophila cretica.

Stem more or less densely glandular-pubescent throughout. Calyx 3-5-5 mm.
long ; costae more or less 3-veined, the lateral veins often very weak. Petals
5°5-7°5 mm. long.

GREECE: Peloponnisos: Androuvista, Montis Taygeti, June-July 1844,
Heldreich (BM ; CGE; K>; isotypes) ; July 1870, Orphanides 593 (W-HAL).
Megala Zonaria, 9 July 1899, Heldreich 1519 (E ; K ; W-HAL). Likoneri, 9 July
1902, Haldcsy 39 (W-HAL).

This taxon has had a somewhat disturbed history. It was first distributed by
Heldreich as Tunica cretica but Boissier described it as a new variety of T. alyrica.

138 REVISION OF THE GENUS PETRORHAGIA

Halacsy, Conspectus Florae Graecae, again considered this plant to be identical with
T. cretica, and appears to have been followed by later authors. Recently Davis
showed that the name 7°. cretica had been misapplied by all authors since Fischer &
Meyer, and that the Cretan plant required a new name. Unfortunately, Davis
followed the majority of authors in regarding Taygetean plants as being identical with
Cretan plants.

We cannot agree with Heldreich, Halacsy or later authors that these two popula-
tions are identical or even that they should be referred to the same species. If the
venation of the calyx is ignored, the Cretan plant still differs from P. zllyrica and the
Taygetean populations by a number of characters and appears to be one of the more
distinct species in this critical group (for further discussion see under P. candica,
p. 141). Here the Taygetean population is regarded as a subspecies of P. illyrica,
showing some affinity with P. candica (cf. the smaller flowers, the feebly 3-veined
calyx costae, and the longer pedicels) but clearly distinct from this latter species.

Subsp. taygetea forms a fairly distinct unit within P. illyrica. It is most readily
recognized by the venation of the calyx costae but similar venation occurs in plants
of typical P. iallyrica in Attiki. The other characters are statistical rather than
absolute differences, so these Taygetean plants are considered to represent a
subspecies rather than a separate species.

2. Petrorhagia ochroleuca (Smith) Ball & Heywood, comb. nov. (Fig. 4.)

Gypsophila ochroleuca Smith, Fl. Graec. Prodr. 1 : 281 (1809).
Tunica ochroleuca (Smith) Fisch. & Meyer, Index Sem. Hort. Bot. Petrop. 4 : 49 (1837).—
Boiss., Fl. Or. 1 : 520 (1867).—Hayek, Prodr. Fl. Penins. Balcan. 1 : 222 (1924).

Stem up to 30 cm. high, usually much branched, glabrous except for the lowest
internodes which are shortly and densely glandular-pubescent, sometimes the
middle and upper parts of the stem sparsely glandular-pubescent (plants from
Evvoia). Lower leaves 10-25 mm. long, linear, 3-veined. Inflorescence lax ;
pedicels 4-14 mm. long. Calyx 4:5-5-5 mm. long, glabrous or rarely sparsely
glandular ; costae 1-veined, sometimes with very feeble lateral veins ; teeth acute
or acuminate. Petals 5~7 mm. long, linear-oblong, entire, pale-yellow, purplish at
the base. Anthers white. Seeds 1-8-2 mm. long, 0-g-1:1 mm. broad, margin not
thickened.

Distribution : South-eastern Greece.

GREECE: Attiki: Hymettus, Heldreich (BM; E; K ; W-HAL); Spruner
(K) ; Haldcsy (E ; W-HAL). Hymettus, versus Pankrati, 13-14 June 1932,
Rechinger fil. 1855 (W-RECH). Parnes, 18 June 1878, Heldreich (W-HAL) ;
18 July 1911, Tunta 1278 (W-HAL). Parnes, Karavola, 29 June 1930, Guiol 1271
(BM). Evvoia: Inter Psachna et Achmet Aga, 27 May 1955, Rechinger fil. 16509
(W). Limni, 28 May 1955, Rechinger fil. 16675 (W) (ad P. armerioidem trans.).

P. ochroleuca is very similar to P. illyrica and is only doubtfully retained as a
distinct species. It can be recognized by a combination of characters (indumentum,
flower and anther colour, pedicel length), none of which on its own serves reliably to

REVISION OF THE GENUS PETRORHAGIA 139

distinguish this species from P. illyrica. The material collected recently by
Rechinger from Evvoia serves only to emphasize the doubtful status of P. ochroleuca.

3. Petrorhagia armerioides (Ser.) Ball & Heywood, comb. nov.

Gypsophila armerioides Ser. in DC., Prodr. 1 : 353 (1824).

Tunica sibthorpii Boiss., Diagn. Pl. Or. Nov. 2 (8) : 61 (1849), nom tllegit. ; Fl. Or. 1: 521
1867).

Pi ares (Ser.) Halacsy, Consp. Fl. Graec. 1 : 194 (1900).—Hayek, Prodr. FI.
Penins. Balcan. 1 : 222 (1924).

Stem up to 30 cm. high, densely glandular-pubescent at the base and apex, usually
glabrous in the middle, but sometimes glandular-pubescent throughout (plants from
Greek mainland). Lower leaves 15-30 mm. long, linear or linear-oblong, 3-veined.
Inflorescence fastigiate or lax with pedicels 1-5(—9) mm. long. Calyx 4-6(—6-5) mm.
long, densely glandular-pubescent ; costae strongly 3-veined ; teeth triangular-
mucronate or acuminate. Petals 6-8(-9) mm. long, oblong-spathulate, white,
often purple-spotted at the base. Anthers purplish. Seeds 1-8-2-1 mm. long,
I-I:3 mm. broad, margin not thickened.

Distribution : South-eastern Greece, Aegean Islands, ? north-western Turkey.

3a. Petrorhagia armerioides var. armerioides. (Fig. 5.)
Tunica argentii Meikle in Kew Bull. 9 : 106 (1954) pro parte, quoad pl. penins. Graec.
Inflorescence dense, fastigiate ; pedicels usually 1-5 mm. long.

GREECE : Sterea Ellas : Graecia, 1836, Zuccarint 592 (K). Ad ruinas Delphorum,
4 Aug. 1904, Maire 41 (W-HAL) (ad P. illyricam trans.)._ Prope Palaeo Kuntura,
montis Pateras, June 1876, Heldreich (W-HAL). Circa Marathon, 1-4 July ro11,
Tunta 1419 (W-HAL). Prope Athena, Reliquiae Orphanideae (E) (ad P. tllyricam
trans.). Aegaleos prope Daphni, 23 June 1905, Bretzl (W-HAL). Parnassos,
supra Arachova, 31 July 1852, Heldreich 5 (BM) (ad. P. allyricam trans.). Mt. Asine,
Argolis, June 1931, Atchley 1103 (K). Prope Vraona, 1 June 1930, Gutol 1468
(BM). Euboea, Mt. Dirphys, 10-17 July 1880, Heldreich (W-HAL) (ad P. allyricam
trans.). Kikladhes : Amorgos, 22-23 May 1808, Dorfler 8,169 (BM ; K ; W-HAL).
Amorgos, ad portam Katapola, 30 June-6 July 1932, Rechinger fil. (W-RECH).
Amorgos, Khozoviotissa, 4 Sept. 1939, Davis 935 (E; K). Naxos, 24-27 June
1889, Heldreich & Halécsy (W-HAL) ; Apr. 1898, Dérfler, Herb. Norm. 3806 (BM ;
E ; W-HAL). Naxos, prope Apiranthos, 16 June 1898, Dérfler 148 (W-HAL).
Naxos, m. Phanariotissa, 28 June 1932, Rechinger fil. 2259 (W-RECH). Tinos
(Tenos), 20 May 1851, Heldreich (K). Tinos, Esomeria, 28 July 1go1, Heldreich
(W-HAL).

30. Petrorhagia armerioides var. laxa (Bornm. & Rech.) Ball & Heywood,

comb. nov.

Tunica armerioides var. laxa Bornm. & Rech. in Beih. Bot. Centralbl. 54 (B) : 614 (1936).
Tunica argentii Meikle in Kew Bull. 9 : 106 (1954), quoad typum, excl. pl. penins. Graec.

Inflorescence lax ; pedicels 3-9 mm. long.
GREECE: Samos: Monte Ambelos, 16-23 June 1932, Rechinger fil. 2098

140 REVISION OF THE GENUS PETRORHAGIA

(W-RECH). Chios: Monte Plaka supra Karies, 12-14 May 1934, Rechinger fil.

5415 (W-RECH). Pelinaion, 12 July 1939, Platt 349 (K, holotype of Tunica
argentit).

Fics. 5-8. 5, Petrorhagia armeriotdes (Ser.) Ball & Heywood var. armerioides. 6, P.candica
Ball & Heywood. 7, P. lycica (P. H. Davis) Ball & Heywood. 8, P. cretica (L.) Ball
& Heywood. a, calyx ; b, petal. (All x5.)

P. armervioides is quite distinct in the Aegean Islands, being distinguished by the
3-veined calyx costae, the indumentum, and the usually short pedicels and dense
inflorescence. However, on the mainland of Greece and Evvoia intermediates
between this species and P. ilyrica or P. ochroleuca, or between all three species, seem
to occur fairly frequently. Nearly all the plants from these areas are to some extent
intermediate as they are usually pubescent throughout the length of the stem.
Plants with lax inflorescence and with feeble lateral veins on the calyx costae also
occur occasionally (e.g. prope Athena, Reliqguiae Orphanideae). It seems desirable
that these populations should be studied in considerable detail, but this must await
further extensive collections particularly in Evvoia and Andros and other islands off

REVISION OF THE GENUS PETRORHAGIA 141

the coast of Attiki. When such material is available it may be found necessary to
regard P. armerioides and P. ochroleuca as subspecies of P. illyrica.

The variety which occurs in Chios and Samos is rather unusual, although it
appears to differ from typical Aegean plants only in the lax inflorescence. It should
not be confused with the long-pedicelled plants occurring on the mainland of Greece
and Evvoia which are intermediates between this species and P. ochroleuca or
P. illyrnica.

4. Petrorhagia candica Ball & Heywood, sp. nov.; inflorescentia laxa pauciflora
absque bracteis epicalycinis, calycis costis 3-venosis, petalis emarginatis retussive
albis subtus roseo-venosis, seminibus nigris laevibus c. 1-5 mm. longis, facile
distinguitur. (Fig. 6.)

Gypsophila cretica Smith, Fl. Graec. Prodr. 1 : 280 (1809) pro parte, quoad descr., excl.
syn..; Pi. Graec:.4 : 76, t. 384 (1823).

Tumica cretica Fisch. & Meyer, Index Sem. Hort. Bot. Petrop. 4: 49 (1837) pro parte,
quoad descr., excl. syn. L.—Boiss., Fl. Or. I : 520 (1867).

Tunica taygetea P. H. Davis in Not. R. Bot. Gard. Edin. 22 : 165 (1957) pro parte, quoad
syn. TI. creticam et Gypsophilam creticam.

Stem up to 20 cm. high, simple or little branched, glabrous or sparsely glandular-
pubescent. Lower leaves 10-20 mm. long, linear-subulate, 3-veined. Inflorescence
lax, few (1-8)-flowered ; pedicels (4—)6-20 mm. long. Calyx (3—)3°5-4:5 mm. long,
glabrous or sparsely glandular-pubescent ; costae 3-veined, the lateral veins some-
times weak ; teeth acute or obtuse-mucronate. Petals 4:-5~7 mm. long, oblong-
spathulate, retuse or emarginate, white with pink veins on the lower surface. Seeds
(slightly immature) c. 1-5 mm. long.

Distribution : Crete.

CRETE : Crete, c. 1930, Guiol (BM; LIVU). Sitia, May 1846, Heldreich (BM, holo-
type ; CGE; K); Szeber 204 (BM; K). Pyrgiotissa, Kryavrisis, 1 June 1904,
Doérfler 384 (W-RECH). Hag. Vasilis, Preveli, 19 June 1904, Dérfler 632 (W-RECH).
Hierapetra, montes Aphendi Kavusi, inter Schinokapsala et Orinon, 20 May 1942,
Rechinger fil. 13183 (K). Sphakia, Levka Ori, inter Samaria et Potamos, 1942,
Rechinger fil. 13802 (BM).

As recently pointed out by Davis (1957), Saponaria cretica L. is the species
described by Fischer and Meyer as Tunica pachygona and not the plant commonly
called T. cretica.

Unfortunately we cannot agree with Halacsy (Consp. Fl. Graec. 1 : 194 (1900)) or
Davis that Tunica cretica auct. is identical with T. illyrica var. taygetea or that they
can be included within the same species, so it is necessary to treat this Cretan plant
as a new species.

P. candica can be readily recognized by the emarginate or retuse petals, the
3-veined calyx costae, longer pedicels and relatively sparse indumentum. The
cauline leaves tend to be shorter and there are more pairs per unit length than in
other species of this section. This, combined with a lesser degree of branching, gives
the plant a strict appearance not found elsewhere in the section.

142 REVISION OF THE GENUS PETRORHAGIA

With some justification Davis minimizes the importance of 3-veined calyx costae,
but his statement that ‘‘ botanists seem to have been hypnotised into seeing them
[3-veined costae] in 7. cretica auct.” is not correct. Nor is his statement that
Rechinger in Flora Aegaea keys out T. cretica auct. as having I-veined costae. The
lateral veins are sometimes feeble, but never completely absent.

All the material seen from Crete has proved to be P. candica while all “‘ Tunica
cretica ’’’ from other areas is referable to other species. The only exception to this is
Baldacci, Iter Creticum 107, which seems to be P. tllyrica subsp. tlyrica. The plants
from this collection are all rather small and the three sheets seen are all very similar,
so that there seems little likelihood of an error. Further investigation at this
locality is desirable. A second collection by Baldacci, [ter Creticum Alterum 293, is
probably wrongly labelled.

5. Petrorhagia lycica (P. H. Davis) Ball & Heywood, comb. nov. (Fig. 7 ;
Plate 13 B.)
Tunica lycica P. H. Davis in Not. R. Bot. Gard. Edin. 22 : 163 (1957).

Stem 10-20 cm. high, branched in the upper part, densely glandular-pubescent.
Lower leaves 10-15 mm. long, linear to linear-spathulate, 3-veined. Inflorescence
lax, the pedicels 3-18 mm. long. Calyx 5-6-5 mm. long, glabrous or sparsely
glandular-pubescent on the costae ; costae 3-veined ; teeth triangular, acuminate
to acute-mucronate. Petals 7-9 mm. long, oblong-spathulate, white, purple-veined
on the lower surface. Seeds 1-5 mm. long, 0-5 mm. broad, brown, minutely
papillose.

Distribution : Turkey (western Anatolia).

TuRKEY : Mugla, Fethiye, Baba Dag above Akbel yayla, 30 July 1947, Davis
13675 (E, holotype ; K). Minara, 31 July 1947, Davis 13709 (BM ; E ; K).

This recently described species does not seem to be very closely related to any of
the other species in this section. Its seeds are more or less intermediate between the
two types found in the genus, but appear to be most similar to those of P. candica.
In other characters it resembles P. armerioides var. laxa and P. ochroleuca, but even
so it seems to be a quite distinct species. For further discussion see Davis (1957).

Subsect. b. Creticae Ball & Heywood

Petrorhagia subsect. Creticae Ball & Heywood, subsect. nov. ; a subsectione [llyricis
differt habitu annuo, seminibus majoribus 2-3-5 mm. longis.
Type : P. cretica (L.) Ball & Heywood.

Annual. Seeds 2—3:5 mm. long.

6. Petrorhagia cretica (L.) Ball & Heywood, comb. nov. (Fig. 8.)

Saponaria cretica L., Sp. Pl., ed. 2, 1: 584 (1762).

Gypsophila cretica (L.) Smith, Fl. Graec. Prodr. 1 : 280 (1809) quoad syn.

Tunica cretica (L.) Fisch. & Meyer, Index Sem. Hort. Bot. Petrop. 4: 49 (1837) quoad
syn. L.

Tunica pachygona Fisch. & Meyer, tom. cit. : 50 (1837).—Boiss., Fl. Or. 1 : 522 (1867).—
Hayek, Prodr. Fl. Penins. Balcan. 1 : 221 (1924).

Gypsophila pachygona (Fisch. & Meyer) D. Dietr., Synops. Pl. 2 : 1543 (1840).

Tunica brachypetala Jaub. & Spach, Ill. Pl. Or. 1: 11 (1842).

REVISION OF THE GENUS PETRORHAGIA 143

Stem up to 40 cm. high, much branched, densely glandular-pubescent, viscid.
Lower leaves 8-25 mm. long, oblong, subobtuse, more or less 3-veined. Inflorescence
lax ; pedicels 4:5—20(—30) mm. long, glabrous or rarely glandular-pubescent. Calyx
6-10°5 mm. long, glabrous ; intercostal membranes 0-g-1:4 mm. broad. Petals
included in the calyx or rarely shortly exserted, linear-spathulate, entire, white,
sometimes reddish on the lower surface. Seeds 2-2-8 mm. long, I-3-I-9 mm. broad.

Distribution : South-western Asia, Greece.

GREECE : Thessalia : Kalampaka, 27 July 1896, Sintents 1223 (E ; K).

TuRKEY : Olympi Bithyn., Pichler 135 (W-HAL). Yaila de Bozdagh (Tmolus
occid.), 17 July 1854, Balansa ro8 (BM ; K). Vallis Meandri, June 1842, Boisster
(BM ; K). Lydia, Magnesia, Mt. Sipylos, g-10 June 1906, Bornmiiller g122 (K).
Elmalu, 13 June 1860, Bourgeau, Pl. Lycide 42 (E; K). Vil. Ankara, Beynam,
5 July 1947, Davis 1306 (BM; E; K). Vil. Ankara, Kibris gorge near Kayas,
6 July 1947, Davis 13141 (BM; E; K). Phrygia, Akscheher, 28 June 1899,
Bornmiiller 4145 (E ; K). N.d’Ouchak, Phrygie vers, 26 July 1857, Balansa 1308
(BM ; K) ; Relig. Matlleanae 224 (BM; K ; W-HAL). Amasia, 24 June 1880,
Bornmiiller r90 (BM ; K). Prov. Adana, district Feke, Gdksu gorge below Himmetli,
g July 1952, Davis 19843 (K). Prov. Maras, district Goksun, Binboga dag above
Yalak, 14 July 1952, Davis 19952 (K). Ad Angora Galatiae, 1892, Bornmiiller 3012
(BM ; E; K). Cappad. orient., Aucher-Eloy 638 (BM ; K). Sert. austr., Aucher-
Eloy 4228 (BM). Armeria turc., Egu, Ekrekdere, 23 June 1890, Sintenis 2682 (K).
Kurdistan 5 km. O. von Siirt, 12 June 1936, Frédin 30 (W). Mardin, 21 June 1888,
Sintenis 1159 (K). Aintab, 16 June 1865, Haussknecht (BM). Inter Aintab et
Marash, June 1907, Haradjian 1493 (W). Mt. Duluk Baba au N. d’Aintab, June
1907, Haradjian 1378 (K).

SyRIA : Supra Damascum, mont. Dschebel Kasium, 13-15 May 1910, Bornmiiller
11459 (BM). Aleppo, Syria-Palestine, Montbret (K).

Cyprus : Between Platres and Aphamis, 16 May 1941, Davis 3445 (E; K\).
Circa Prodromo, in mt. Trodos, 17 June 1880, Sintenis & Rigo g60 (CGE ; K).
Platres, 9g June, 18 June, 27 June 1938, Kennedy (K). Near Philani, 13 May 1908,
Clement Reid (K). Parakklisha-Kellaki, Limasol, 24 May 1948, Kennedy 1615 (Kx).

IRAQ : Gabal Hamrin, N.E. of Baghdad, 24 Nov. 1918 (seed, hort., Aug. 1920),
W. E. Evans M/3 (E). Mam Zawita district (Lomana), 1-8 June 1955, Robertson
14456 (K). Jarmo, 1 May 1955, Helbaek 1286 (K). Jebel Khatchra near Balad
Sinjar, 5 June 1934, Field & Lazar 646 (KK). Pl. Mesopot., Kurdistan et Mossul,
1841, Kotschy 331 (K).

IRAN : Azna, 12 June 1937, Kéte 1378 (W).

For discussion of the nomenclature of this species see Davis (1957). The type
of Saponaria cretica is undoubtedly this species (see p. 141).

7. Petrorhagia arabica (Boiss.) Ball & Heywood, comb. nov. (Fig. 9.)

Tunica arabica Boiss., Diagn. Pl. Or. Nov. 2 (8) : 62 (1849) ; Fl. Or. 1 : 523 (1867).
Tunica pachygona var. hirtituba F. N. Williams in Journ. of Bot. 28 : 199 (1890).

144 REVISION OF THE GENUS PETRORHAGIA

Stem up to 30 cm. high, much branched, glandular-pubescent, viscid. Lower
leaves 15-30 mm. long, linear-oblong, subobtuse, more or less 3-veined. In-
florescence lax, with pedicels 3-30 mm. long, densely glandular-pubescent. Calyx
6-5-9 mm. long, densely glandular-pubescent ; intercostal membranes 0-7—1-I mm.
broad. Petals 9-12 mm. long, distinctly longer than the sepals, linear-spathulate,
entire, white. Seeds 2-7—-3:5 mm. long, 1-6-2 mm. broad.

[2

Fics. 9-12. 9, Petrorhagia arabica (Boiss.) Ball & Heywood. 10, P. hispidula (Boiss. &
Heldr.) Ball & Heywood. 11, P. alpina (Habl.) Ball & Heywood subsp. alpina. 12,
P. kennedyae (Jacks. & Turrill) Ball & Heywood. a, calyx ; b, petal. (All x5.)

Distribution : Israel and Jordan.

ISRAEL, JORDAN : Magdala, 1863-64, Lowne (BM ; CGE; E; K). Mt. Gilboa,
top of pass from Bari to Faggira, 17 May 1942, Davis 4739 (E ; K). East of Jordan,
1873, J. A. Paine (K). Wad-el-Kat, 31 Mar. 1909, Meyers & Dinsmore 5311
(E ; K). South of Shahba, 28 June 1932, Dinsmore rrorz (K). N. of Nablus,
I June 1935, Eig, Feinbrun & Leinkram 320 (E ; K). Inter Tiberiadem et Safed,
9 June 1881, Letourneux 328 (K). Shittim plain, 24 Apr. 1886, Post (K).

REVISION OF THE GENUS PETRORHAGIA 145

This species generally appears to be distinct from P. cretica. It can be dis-
tinguished by the usually much larger seeds, the pubescent calyx with compara-
tively narrow intercostal membranes, and the petals always distinctly longer than
the sepals. PP. cretica rarely has the calyx very slightly glandular at the base and
more frequently has the petals slightly longer than the sepals. The frequency
with which these apparent intermediates occur is uncertain, but if they should
prove to be widespread in Syria, Israel and Jordan then P. arabica should perhaps
be regarded as a subspecies of P. cretica. However, from the sparse material seen
these two species appear to be distinct in this area, although there has undoubtedly
been some confusion between them.

8. Petrorhagia hispidula (Boiss. & Heldr.) Ball & Heywood, comb. nov. (Fig.10.)
Tunica hispidula Boiss. & Heldr. in Boiss., Diagn. Pl. Or. Nov. 2 (8) : 62 (1849).—Boiss.,
POP S 5231867).

Stem up to 30 cm. high, much branched, glandular-pubescent, viscid. Lower
leaves 10-25 mm. long, linear-oblong, linear or linear-subulate, more or less 3-
veined. Inflorescence lax, with pedicels 4-15 mm. long, densely glandular-pubescent.
Calyx 5-5-7 mm. long, pubescent or subglabrous ; intercostal membranes 0-4—0-7
mm. broad. Petals 5-5-8 mm. long, included in the calyx or slightly exserted,
linear-spathulate, 4-lobed, white. Seeds 1-8—2:2 mm. long, 1-2-1-7 mm. broad.

Distribution : Turkey (southern Anatolia).
TuRKEY : Inter Adalia et Maela, July 1845, Heldreich (E ; K; W; W-HAL ;

isotypes). Supra Adalia, July 1845, Heldreich (E). Near Gebiz, 22 July 1949,
Davis 15476 (BM ; E; K).

Sect. 2. PSEUDOGYPSOPHILA (A. Braun) Ball & Heywood

Petrorhagia sect. Pseudogypsophila (A. Braun) Ball & Heywood, comb. nov.

Tunica subgen. Leptopleura Jaub. & Spach, Ill. Pl. Or. 1: 11 (1842).
Tunica sect. Pseudogypsophila A. Braun in Flora 26 : 384 (1843).
Tunica sect. Leptopleura (Jaub. & Spach) Boiss., Fl. Or. 1 : 521 (1867).

Type : Tunica stricta (Ledeb.) Fisch. & Meyer (=P. alpina (Habl.) Ball &
Heywood).

Annual. Leaves 1-5-veined. Epicalyx bracts absent. Petals not clawed.
Seeds small, blackish-brown, reticulate-tuberculate, with thickened margin.

9. Petrorhagia alpina (Habl.) Ball & Heywood, comb. nov.

Gypsophila alpina Habl., Neue Nord. Beitr. 4 : 57 (1783) ; in S. G. Gmel., Reise Russl. 4 :
178 (1784).

Gypsophila stricta Ledeb., Ic. Pl. 1 : 4 (1829).—Bunge in Ledeb., Fl. Alt. 2 : 129 (1830).

Tunica stricta (Ledeb.) Fisch. & Meyer, Index Sem. Hort. Bot. Petrop. 4: 50 (1837).—
Boiss:, Bl, Or: 1..> .522 (1867).

Dianthus recticaulis Fenzl in Ledeb., Fl. Ross. 1 : 287 (1842).

Tunica alpina (Habl.) Bobrov, Bot. Zhurn. 43 : 1546 (1958).

Stem up to 40 cm. high, erect, much branched, glabrous. Basal leaves rosulate,

146 REVISION OF THE GENUS PETRORHAGIA

5-30 mm. long, 1-4(-7) mm. broad, oblong- to linear-spathulate, acute or obtuse ;
cauline leaves linear, acute, I-veined. Calyx 2-5-4:5(-5:5) mm. long, glabrous,
green ; teeth shortly triangular-acute or obtuse-mucronate. Petals (3—-)3-5-6 mm.
long, linear-oblong, entire, white. Seeds 0-7-1-2 mm. long, 0-4-0-7(-0-9) mm.
broad, ovate-oblong.

Distribution : Mountains of western and central Asia, eastwards to the western
Himalaya ; southern Bulgaria.

ga. Petrorhagia alpina subsp. alpina. (Fig. 11.)

Leaves oblong-spathulate. Inflorescence strict ; pedicels 0-5—12(-20) mm. long,
erect.

TuRKEY : Nigde, Hasan Dag below Taspinar Yayla, 17 June 1952, Davis 19003
(BM; E; K). Rize, Vallée de Cimil, 16 July 1866, Balansa 1415 (BM; K).
Kayseri, Bakir Dag, above Kisge, 2,000 m., 30 June 1952, Davis 19385 (BM ;
E ; K). Adradices mont. Tech-Dagh, supra Erzeroum, July 1853, Huet (BM ; K).
Giresun, Balaban daglari above Tamdere, 1,800 m., 6 Aug. 1952, Davis 20510
(BM; E; K) (ad subsp. olympicam vergens). Van, Catak, Kavussahap Dag,
2,900 m., 24 July 1954, Davis 23063 (BM; E; K). (Coruh, Kordevan dag near
Kutul yayla, 28 June 1957, Davis 30237 (E). Kars, Yalnizcam Daglari above
Yalnizgam, 2,100-2,300 m., 19 Aug. 1957, Davis 32501 (E). Erzincan, Kesis dag
above Cimin, 2,300 m., 28 July 1957, Davis 31832B (E).

U.S.S.R.: Armenia, Migri, inter et supra Vargavar et Lishkras, I,500-1,600 m.,
29 June 1934, Karjagin (BM). Ahas Tauman, Iber. occid., Smirnoff (W-HAL).
Azerbaidjan, Nakhichevanskaya, 17 June 1947, Grossheim, Ilinskaya & Kirpichnikov
(BM). Kazakhstan, Semipalatin, Karkaraly, 1890, Korjinsky (BM). Altai,
Ledebour (K). Dschany-bek, mont. Tarbagatai, mont. Kurtschum, 1840, Karelin
& Kiriloff 136 (BM ; K). Prope Kolyvanskoje Osero, 12 May 1go1, Krylov (K).
Tadzhikistan, Iovon, 10 June 1893, Komarov (K). Tadzhikistan, Kuli-Mohif,
May 1893, Komarov (K). .

IRAN : Mazanderan, in valle fluvii Calus, 9 June 1937, Rechinger fil. 913 (BM ;
K). Tochtal, 3,300 m., July 1935, Lindsay 511 (BM). M. Demaurend, 2,100 m.,
21 June 1843, Kotschy 354 (BM ; K). Alatau ad fl. Lepys, Kotschy 1296 (BM).

PAKISTAN, KASHMIR: Kashmir, Falconer 237 (K). Huripur to Aliabad,
Falconer 2984 (K). Shapujon, 2,100 m., 9 July 1876, Clarke 28596 (K). Aliabad,
2,700 m., 9 July 1876, Clarke 28677 (K). Nowgunje, 2,000 m., 19 July 1876,
Clarke 29199 (K). Kunshoan, 2,250 m., Clarke 29368 (K). Liddar, S. Kashmir,
2,650 m., 5 July 1902, Drummond 14128 (K). Between Aru and Lidarwat, 6 July
1902, Drummond 14125 (K). Ushkra, 1,600 m., July 1891, McDruell (K). Kammi
pass, above Shankangarh, 3,000 m., 16 July 1946, Stewart 22764 (K). Near Gulmarg,
2,400-2,700 m., 19 June 1893, Duthie 13016 (BM). Swat, Utrot, 2,400 m., Stewart
& Rahman 25269 (BM).

gb. Petrorhagia alpina subsp. olympica (Boiss.) Ball & Heywood, stat. nov.
(Plate 13 C.)

REVISION OF THE GENUS PETRORHAGIA 147

Tunica olympica Boiss., Diagn. Pl. Or. Nov. 2 (8) : 61 (1849) ; Fl. Or. 1: 522 (1867).
Tunica filiformis Post in Bull. Herb. Boiss. 1 : 411 (1893).
Leaves linear-spathulate. Inflorescence spreading ; pedicels (0:5—-)3-30 mm.
long, patent.

BuLGaRiA : In pinetis m. Pirin ad Suchodol, 14 July 1929, Stojanoff, Stefanoff
& Georgieff (K).

TURKEY : Olympus, Aug. 1842, Boissier (BM ; E; K ; isotypes) ; June-July
1862, Mill (BM ; K) ; July 1874, Pichler 135 (K) ; Bornmiiller 446 (E). Uludag,
1,800-2,180 m., 19 June 1956, Moore 7287 (E). M. Ida, prope Kareikas, 1893,
Sintenis 599 (BM ; E ; K ; W-HAL). Kutahya, Gediz, Saphane dag, 6 Aug. 1950,
Davis 18463 (BM ; E; K). Kutahya, Vili Muphlor, Sundras dag at Goékce ova,
27 July 1947, Davis 13613 (E). Vergers de l’Yaila de Bozdagh (Izmir), 28 July
1854, Balansa 105 (BM; K). Phrygia, Sultan dagh, Akscher, 25 June 1899,
Bornmiiller 4147 (E ; K). Isparta, Sutculer (Isauria), between Tota bebi yayla and
Daribiiykii, 30 July 1949, Davis 15882 (BM; E; K). Isparta, Sutculer, Dedegol
dag, between Selkose and Oruz Gazi, yayla, 1,300 m., 1 Aug. 1949, Davis 15929
(E). Isparta, Sutculer, Dedegdl dag between Daribiiykii and Selkose, 30 July
1949, Davis 15866 (E; K). Wilajet Kastambuli (Kastamonu), Tossia in pinetis
ad Lhakirla (Cakirlar), 16 June 1892, Sintenis 4651 (BM). Kure-Nahas (near
Inebolu), 5 Sept. 1892, Sintenis 5150 (BM ; K). Amasia, m. Ak-dagh, 18 June
1889, Bornmiiller 184 (BM ; K) (ad subsp. alpinam vergens). Cappad. orient.,
Aucher-Eloy 556 (BM ; K) ; Montbret (K). Taurus, au nord du defile des Portes
cilicieuses (Gulek Bogaz), June-July 1855, Balansa (EF). Anatolia orientalis,
Sandz, Liggna-dagh, 4 July 1917, Schischkin (BM). Mt. Amanus, July 1801,
Post (K, isotype of Tunica filiformis) ; 1906, Haradjian 381 (K) ; 800-1,000 m.,
Sept. 1913, Haradjian 4685 (W). Kusliji dagh, 1,700-2,200 m., Aug. 1908,
Haradjian 2532 (K ; W).

Bobrov (1958) has recently shown that Gypsophila alpina Habl. is identical
with Tunica stricta (Ledeb.) Fisch. & Meyer. Hablizl’s epithet is therefore the
earliest for the species.

Boissier listed a number of characters by which Tunica olympica could be dis-
tinguished from 7. stvicta (spreading panicle, longer slender pedicels, smaller obovate
seeds). Our investigations have shown that these characters are not very satis-
factory. The seeds of both taxa appear to be identical, while there is such a
wide range in pedicel length, even on the same plant, that this can only be used on
a statistical basis. The difference in the inflorescence, although reasonably satis-
factory, is not always reliable, and there seems to be some intergradation in northern,
central and eastern Turkey. We therefore treat T. olympica as a subspecies of
P. alpina.

10. Petrorhagia kennedyae (Jacks. & Turrill) Ball & Heywood, comb. nov.
(Fig. 12 ; Plate 13 p.)
Tunica kennedyae Jacks. & Turrill in Bull. Misc. Inf. Kew 1938 : 462 (1938).
Tunica cypria Rech. f. in Fedde, Repert. Sp. Nov. 47 : 163 (1939).

148 REVISION OF THE GENUS PETRORHAGIA

Stem 5-30 cm. high, erect, usually branched from the base, shortly glandular-
hispid at the base, glabrous or glabrescent in the middle, glandular and viscid at
the apex. Basal leaves up to 12 mm. long and 4 mm. broad, oblong-elliptic,
obtuse ; cauline leaves 5-15 mm. long, 0-5-2 mm. broad, 3-veined, linear, acute.
Inflorescence lax, many-flowered ; pedicels 5-25 mm. long, spreading. Calyx
4°5-6:5 mm. long, glandular-viscid, the costae often reddish or purplish ; teeth
broadly triangular-acute or obtuse-mucronate. Petals 6-8-5 mm. long, linear-
spathulate, white, usually reddish on the lower surface, bifid. Seeds 1:3-1-6 mm.
long, 0-6—0-8 mm. broad, oblong.

Distribution : Cyprus.

Cyprus : Troodos mountains, 17 July 1880, Sintenis & Rigo 764 (BM ; CGE ;
K ; W-HAL) ; 20-25 June 1912, Haradjian 467 (K). Platres, 18 June 1938,
Kennedy 1037 (K, holotype). Mesopotamus, 21 June 1939, Lindburg (K) ; on
dry igneous slopes in Pinus forest, 1,200 m., 16 May 1941, Davis 3450 (E ; K).
Krigos, Potamos, 20 June 1938, Kennedy 1038 (K). Stevros (Paphos), 1,200 m.,
3 July 1940, Davis 1759 (E ; K). Rondhkias valley between Pano and Kykko,
450-600 m., 8 May 1941, Davis 3385 (E ; K). Above Alithenon, Merton 2408 (K).

This species appears to have been first collected as long ago as 1880 by Sintenis
and Rigo, but was not recognized until 1938 when Jackson & Turrill published a
description ; soon afterwards Rechinger described it independently.

The relationships of this species are somewhat uncertain. Jackson & Turrill
placed it in Tunica sect. Pachypleura, primarily because of its superficial resemblance
to P. cretica (Tunica pachygona). It resembles that species in its annual habit,
glandular indumentum and comparatively large flowers, but differs in the
one-veined calyx costae and the small, blackish-brown scutate seeds. These differ-
ences are here considered to be of major taxonomic significance and indicate that
this species should be placed in Sect. Pseudogypsophila together with P. alpina,
which is also an annual species. However, P. kennedyae is quite distinct from
P. alpina, and can be distinguished by a series of characters (e.g. indumentum,
larger flowers, bifid petals, etc.).

Sect. 3. PETRORHAGIA

Petrorhagia sect. Petrorhagia ; sectio typica generis.

Tunica sect. Gypsophiloides Griseb., Spicil. Fl. Rumel. 1 : 184 (1843), nom. illegit.
Tunica sect. Pseudodianthus A. Braun in Flora 26 : 384 (1843), nom. illegit.

Tunica sect. Eutunica Boiss., Fl. Or. 1 : 518 (1867), nom. illegit.

Tunica sect. Tunicastrum F. N. Williams in Journ. of Bot. 28 : 194 (1890), nom. illegit.

Type: P. saxifraga (L.) Link.

Perennial. Leaves I-veined (rarely up to 5-veined). Flowers with or without
epicalyx bracts. Petals not abruptly clawed. Seeds blackish-brown, tuberculate,
with thickened margin.

Subsect. a. Saxifragae Ball & Heywood
Petrorhagia subsect. Saxifragae Ball & Heywood, subsect. nov.; subsectio typica

REVISION OF THE GENUS PETRORHAGIA 149

sectionis Petrorhagiae, continens species caulibus valde ramosis, floribus solitariis
vel fasciculatis, bracteis lanceolatis vel ovatis.
Type: P. saxifraga (L.) Link.
Stems much branched. Flowers solitary or fasciculate ; bracts and epicalyx
bracts (when present) lanceolate or ovate, membranous, I-veined.

11. Petrorhagia phthiotica (Boiss. & Heldr.) Ball & Heywood, comb. nov.
(Fig. 13 ; Plate 14 A.)
Tunica phthiotica Boiss. & Heldr. in Boiss., Fl. Or., Suppl. : 82 (1888).
Tunica ochroleuca var. phthiotica (Boiss. & Heldr.) Hayek, Prodr. Fl. Penins. Balcan. 1 :
223 (1924).

Stems 3-20 cm. high, sparsely glandular-pubescent or subglabrous. Lower
leaves 3-8 mm. long, I-3-veined. Inflorescence lax ; epicalyx absent. Calyx
3-5 mm. long, glabrous ; costae I-veined ; teeth broadly triangular to very obtuse,
mucronate. Petals 4-6-5 mm. long, 0-5-1 mm. broad, oblong-spathulate, entire,
pink. Seeds 1-2—1-6 mm. long, 0:55—0-75 mm. broad.

[5 16

Fics. 13-16. 13, Petrorhagia phthiotica (Boiss. & Heldr.) Ball & Heywood. 14, P. fascicu-
lata (Marg. & Reut.) Ball & Heywood. 15, P. saxifraga (L.) Link var. saxifraga. 16,
P. graminea (Smith) Ball & Heywood. a, calyx ; b, petal ; c, epicalyx bract. (All x 5.)

Distribution : South-eastern Greece (Oiti) ; very doubtful in north Peloponnisos.

GREECE : Sterea Ellas: M. Oeta, 14 July 1879, ?Heldreich (W-HAL, ?isotype) ;
30 June 1935, Guiol 2366 (BM). Katafigon, Oeta, 11 June 1937, Balls & Gourlay
3244 (BM ; K). Peloponnisos : Kyllene at head of the gorge of Phlambouritza,
June 1862, Mill (K) (? locality correct).

150 REVISION OF THE GENUS PETRORHAGIA

The relationships of this species are somewhat uncertain, although it is perhaps
nearest to P. fasciculata. However, P. phthiotica also shows some resemblance to
P. alpina and it may represent a link between Sects. Petrorhagia and Pseudogypsophila.
It was originally placed in his Tunica sect. Gypsophiloides by Boissier, but the very
different seed type, in addition to a number of secondary characters, clearly dis-
tinguishes it from the other members of that group, which we place under Petrorhagia
sect. Pseudotunica. It is difficult to understand why Hayek regarded it as only a
variety of Tunica ochroleuca, but this may have been due to the paucity of material,
and the failure to recognize the importance of the seed characters. Apart from the
seeds, P. phthiotica can easily be distinguished from P. ochroleuca by the smaller
pink or purplish flowers, the broad triangular mucronate calyx teeth, and the different
indumentum (cf. Davis, 1957).

P. phthiotica differs from other members of Sect. Petrorhagia in a number of
characters, the significance of which is somewhat uncertain. The distinctive
calyx teeth, glabrous calyx, entire petals, complete absence of fasciculation of
flowers, and the sparse patent-glandular hairs are the more obvious differences,
and these may well prove to be more indicative of relationships than the perennial
habit, the very narrow leaves, and the pink flowers.

12. Petrorhagia fasciculata (Marg. & Reut.) Ball & Heywood, comb. nov.
(Fig. 14.)
Gypsophila fasciculata Marg. & Reut. in Mém. Soc. Phys. & Hist. Nat. Genéve 8: 281
(1839).—Boiss., Fl. Or. 1 : 555 (1867).
Tunica fasciculata (Marg. & Reut.) Boiss., Fl. Or., Suppl. : 82 (1888).—Hayek, Prodr. F1.
Penins. Balcan. 1 : 223 (1924).

Stems 5-30 cm. high, glandular-pubescent, sometimes glabrous above. Lower
leaves 6-30 mm. long, 0-5—-1-8 mm. broad, usually 1-veined, rarely up to 5-veined.
Inflorescence branched ; flowers usually fasciculate and bracteate, rarely solitary
and then ebracteate. Calyx 2:5—5:5 mm. long, shortly or sparsely hirsute or glandu-
lar ; costae I-veined ; teeth acute or subobtuse. Petals 3-5-6 mm. long, linear-
spathulate, retuse, white or pale-yellow. Seeds 0-8-1 mm. long, 0-6—0-7 mm. broad.

Distribution : Western Greece and Ionian Islands.

GREECE : Sterea Ellas : Stratos, Aetolia, 1935, Guiol 2439 (BM). Arcania, prope
Agrinion, 3 July 1893, Haldcsy (K ; W-HAL). Zacynthos, Mazztoiri (W-HAL).
Hag. Ilias, Levkas, 30 June 1895, Baldacci 13 (BM ; K ; W-HAL). Kephallonia,
Same (Lixourion), 1926, Bornmiiller 188 (BM ; K), rg0 (BM ; K). Cephallonia,
mont. Phalaris, Aug. 1867, Heldreich (K).

This species appears to be a reasonably distinct member of Sect. Petrorhagia
and can be recognized by the white or pale-yellow, usually fasciculate flowers, the
smaller calyx With more or less acute teeth and one-veined costae, and the patent,
glandular hairs (sometimes even on the calyx). These last two characters are
shared with P. phthiotica and this to some extent supports the view that P. phthiotica
should be placed in this section. However, P. fasciculata has a number of important
features in common with the other members of the section (e.g. the fasciculation

REVISION OF THE GENUS PETRORHAGIA 151

of the flowers, the white hirtulose calyx indumentum, the more or less oblong
calyx teeth, and the very narrow linear leaves) so that there can be no grounds
for regarding it as separate.

It should be noted that plants of P. fasciculata sometimes occur with all the
flowers solitary, which are then ebracteate.

13. Petrorhagia saxifraga (L.) Link, Handb. Erkenn. Gewachse 2 : 235 (1831).
Dianthus saxifragus L., Sp. Pl. 1 : 413 (1753).
Gypsophila saxifraga (L.) L., Syst. Nat., ed. 10, 2 : 1028 (1759).
Tunica saxifraga (L.) Scop., Fl. Carniol., ed. 2, I : 300 (1772).
Dianthus filiformis Lam., Fl. Frang. 2 : 537 (1778), nom. illegit.
Imperatia filiformis Moench, Meth. Pl. : 60 (1794), nom. tllegit.
Gypsophila multicaulis Poir. in Encycl. Méth., Bot., Suppl. 2 : 875 (1812).
Petrorhagia multicaulis (Poir.) Link, loc. cit. (1831).
Gypsophila arenicola Dufour in Bull. Soc. Bot. France 7 : 240 (1860).
Tunica erecta Jord. & Fourr., Brev. Pl. Nov. 1 : 10 (1866).
Tunica bicolor Jord. & Fourr., loc. cit. (1866).
Tunica xerophila Jord. & Fourr., loc. cit. (1866).
Tunica ciliata Dulac, Fl. Dép. Haut.-Pyrén. : 260 (1867), nom. illegit.
Tunica arenicola (Dufour) F. N. Williams in Journ. of Bot. 28 : 194 (1890).
Kohlrauschia saxifraga (L.) Dandy in Watsonia 4 : 42 (1957).

Stems 5-45 cm. high, glabrous, shortly papillose or scabrid-pubescent. Lower
leaves 5-25 mm. long, 0-4-1°5(—2:5) mm. broad, I-veined. Inflorescence lax or
fasciculate ; flowers usually with 4 epicalyx bracts. Calyx 3-6(-7) mm. long,
glabrous or occasionally sparsely pubescent ; costae I-veined with 2 weak lateral
veins ; teeth oblong, obtuse. Petals 45-10 mm. long, 1-2-3(-4) mm. broad, the
limb obcordate, white or pink. Seeds o-g-1-6 mm. long, 0-6—-1-I mm. broad.

Distribution : Central and southern Europe and south-western Asia.

13a. Petrorhagia saxifraga var. saxifraga. (Fig. 15 ; Plate 14 B.)
Lower leaves 0:4-1:5 mm. broad. Flowers solitary. Calyx 3-6 mm. long.

PorTuUGAL : Porto, 1848-50, Welwitsch 851 (BM; CGE). Carrica, margem
direita do rio Douro a 2 km. de Barca d’Alva, 27 June 1955, Fernandes, Matos &
Matos, Flora Lusitanica 5664 (BM).

SPAIN : Valentia, montis Segaria, pr. Dianium, 8 May 1923, Gros, Flora Iberica 119
(BM ; K). Denia, Alicante, 12 May 1928, Ellman & Sandwith 1132 (K).

FRANCE : Gard: West of river Rhone opposite Avignon, 13 June 1952, Souster
1249 (K). Vaucluse : Serignon, 1 June 1880, Delacour (K). Dréme : Romans-sur-Isére,
June 1872, anon. (W-HAL). Rhdne: Silleuxbanne prés Lyon, 19 June 1859,
Guichard (K ; W-HAL). Ain: Balan, 10 Sept. 1865, Miciol (BM ; W-HAL).
Jura: Sept. 1857, Herb. E. A. Wilmott (K). Haute Savoie: La montagne de
Mandallax prés d’Annecy, Billot 1431(2) (BM ; CGE). Savore : Mont. Lemene,
prés de Chambéry, Billot 1431(r) (BM; CGE). JIsére: Grenoble, 6 July 1884,
Magnier (W-HAL). Hautes-Alpes ; Briancon, Aug. 1870, Reverchon (K). Basses-
Alpes : Le Roc, Castellare, 27 June 1950, Brenan 9781 (K). Var: St. Raphael,

152 REVISION OF THE GENUS PETRORHAGIA

3 June 1873 (FI). Alpes-Maritimes : Menton, Dec. 1866, Joad (E) ; Rechinger fil.
1507 (W-RECH) ; Nov. 1920, Lester-Garland (K).

Corsica : Evisa, 13 June, 28 July 1884, Reverchon (BM ; E ; FI; K ; W-HAL).
Pentes du Pigno, a Bastia, 2-13 June 1867, Mabille 21b (BM ; CGE; E; FI; K).
Vizzavona, July 1907, Martelli (FI). Ragliano, 26 Sept. 1854, Reveliére 43 (BM).

GERMANY: Munchen, Sfitzel 393 (BM; CGE; E; K). Prope Riesenfeld,
1 Aug. 1883, Woerlein (LIVU).

SWITZERLAND : Geneva, June 1893, Ndgeli (CGE ; E) ; Botssier (K). Between
Visp and Zermatt, Evans (E). Lausanne, Aug. 1879, Favrat & Barbey (K). Tarasp,
unter Engadin, Aug. 1903, Crawford (E). Between Sierre and Vissoye, Aug. 1903,
Lester-Garland (K).

Austria : Tirol: Innsbruck, Sept. 1882, Evers (W-RECH). Botzen, 17 Aug.
1860, Babington (K). Ost Tivol: Lienz, June 1871, Gander (E; K; W-HAL).
Karnten : (Klagenfurt, July 1872, Ehrenberger (W-HAL). Stevermark: Graz,
July 1901, Hayek, Fl. Stir. 956 (BM ; E). Ober-Osterreich : Prope Styr, Dérfler,
Fl. Austro-Hung. 3228 I (BM ; E;K ; W-HAL; W-RECH). Nieder-Osterreich :
Marchfeld, 21 June 1910, Korb (W). Wien, Aug. 1876, Haldcsy (W-HAL).
Burgenland : Podersdorf, 24 June 1923, Rechinger (W-RECH).

CZECHOSLOVAKIA : Slovakia austr.-occid., prope Plavecky Ctvrtek, 20 June 1929,
Domin & Krajina 35 (CGE ; K).

Huncary : Fehér, Fehérvarcsurgé, 28 June 1923, Fl. Hung. 836 (BM; E;
K). Tolna, ad stationem Tolnanemeti, 8 July 1922, Pillich (BM ; K). Szontagh,
Odenburg (Sopron), 1861, Sempront (K).

RoMANIA : Verciorova, 5 June 1887, Degen 20 (E; K; W-HAL). Baile
Herculane, June 1907, Schneider 1215 (BM; K). Gura Vaii, June 1881, Grecescu
(W-HAL). Kudijévar, 19 June 1875, Tauscher (E).

ITaLy : Piemonte: Aosta, 16 June 1898, Vaccari (FI). Susa, 8 Aug. 1854,
Parlatore (FI). Lombardia : Bormio, June 1870, Sommier (FI). Liguria: San
Remo, Mar. 1880, Marchesetti (FI). Genova, Aug. 1856, Ardissone (FI). Venezia :
Lungo il Bachiglione presso Padora, 3 June 1894, Fiori (FI). Venezia Carperica
Vittorio, 6 June 1894, Pampanini (FI). Tridentina : S. Leonardo in Passiria,
V. Valtina, 17 July 1949, Zenari (FI). Toscana: Firenze, Sept. 1867, Sept. 1875,
Groves (FI; K). We d’Elbe, 27 Feb. 1871, Sommier (FI). Grossetto, M. Orace,
14 June 1918, Fiori (FI). Siena, Colle V. d’Elsa, 17 June 1918, Fiori (FI). Marche :
Salendo al Vettore (Mti. Sibillini), 10 Aug. 1895, Sommier (FI). Ascoli, 3 July
1856, Parlatore (FI). Emilia: Riola, Rochetta, July 1883, Baldacci (FI). San
Marino, 15 Oct. 1916, Pampanini 1267 (FI). Modena, 20 June 1882, Fiori (FI).
Lazio: Tivoli, M. Rigali, Sept. 1901, Vaccari (FI). Terracina, 22 May 1893,
Sommier (FI). Presso il Colosseo, Roma, 12 May 1886, Aiutz (FI) ; May 1836,
Trevelyan (CGE). Abbruzzo: M. di Caramanica, 6 July 1872, Pedicino (FI).
Majella, in valli Orfente, ad radic. montis Mucchia, 31 July 1874, Levier (FI).
Campania ; Sorrento, June 1911, Guadagno (FI). Monte Alburno, 29 May 1951,

REVISION OF THE GENUS PETRORHAGIA 153

Philippis & Moggi (FI). Capri, May 1843, Heldreich (K). Umbra: Mura di
Assisi, 14 June 1935, Corradi (FI). Ierni, Alle Cascate, 26 June 1875, Marchesetti
(FI). Puglia: Taranto, June 1919, Lacaita, Fl. Ital. 2434 (BM ; FI; K). Monte
Nero presso S. Marco, 10 May 1893, Martelli (Fl). Basilicata: Mt. Sena delli
Ciovoli, 4 Aug. 1929, Gavioli 14996 (FI). Vulture, July 1928, Biondi (FI). Calabria:
Palizzi, 9 June 1877, Arcangeli (FI). Reggio, 15 May 1872, Anztz (FI).

SARDINIA : Sadali, 24 July 1898, Martelli (FI). Terranora Pousania, 16 June
1899, Martelli (FI).

Sicity : Etna, Sept. 1841, Parlatore (FI). Palermo, May 1905, Ross 515 (BM ;
E ; FI). Madonie a Quacedda, June 1840, Parlatore (FI). Ins. Favigna, 5 May
1855, Huet (BM ; CGE; FI; K). Sorrentino, July 1845, Sorrentino (FI).

Jucostavia : Slovenija: Near Bled, 1923, Leathes (BM). Radmannsdorf,
Aug., Paulin, Fl. Carniolica 262 (BM). Hrvatska: Zagreb, 2 July 1879, Rossi
(K). Pola, Capo Promontore, 6 June 1919, Vaccari (FI). Spalato (Split), Pichler,
Fl. Austro-Hung. 3228 II (BM ; E; W-HAL ; W-RECH). Lesina, 18 May rg10,
Keller (W-HAL). Dubrovnik, 26 June 1953, Dennis 52 (K). Srbya: Nischka
Banja prope Nisch, 15 June 1955, Rechinger fil. 16064 (W). Vranja, 17 July 1887,
Bornmiiller (E). Sveto-Petra, 26 Aug. 1921, Yermoloff (BM). Novi Pazar, 8 July
1956, Carpenter 342 (K). Bosna 1 Hercegovina: Nevesinje, 23 Aug. 1895, Callier
m7t (K; W-HAL). Sarajevo, 22 June 1930, Gillrat-Smith 2662 (K). Mont.
Porim, prope Mostar, July 1893, Vandas (K). Crna Gora: Lovcen, 9 July 1927,
Korb (W). Mt. Jerinja, July 1906, Rohlena (BM). In colle Ljinbovic ad Podgorica,
11 Sept. 1902, Baldacci 212 (BM). Kostat, 14 July 1891, Baldacci (E ; W-HAL).
Makedonija : Inter Debar et Mavrovi Hanovi, 12 June 1955, Rechinger fil. 15796
(W). Gjefgjeli, 1-2 June 1927, Rechinger fil. 1564 (W-RECH). Kaunferiji, June
1906, Adamovié (BM).

ALBANIA : Pastrik, 31 Aug. 1916, Dérfler 343 (E; K). West of L. Ohridsko,
13 Aug. 1938, Hepburn 84 (K). Sarandé, 3 June 1933, Alston & Sandwith 1339
(BM ; K). M. Kusema, supra Vonitko, dist. Leskovik, 9 July 1896, Baldacci 215
(K ; W-HAL). Supra Sceperi, dist. Tepelené, 22 July 1894, Baldacci 55 (BM ; E).

ButcarRiaA : Near Rila, 2 Aug. 1926, Turrill 1541b (K) ; 2 Aug. 1926, Stojanoff
& Stefanoff (BM). Causovo, June 1908, Strzbrny (BM).

GREECE : Ipiros: Arta, 5 July 1893, Haldcsy (W-HAL). Mt. Peristeri, 11 July
1893, Haldcsy (W-HAL). Metsovon, June-July 1929, Guiol 1467 (BM). Cephalonia,
19 Oct. 1834, Schimper & Wrest (K). Makedhonia: Prespa L., Io June 1932,
Alston & Sandwith 250 (BM; K). Between Turica and Gumus Dere, south of
Struma Plain, 5 June 1917, Turrill 381 (K). Peloponnisos: In m. Omplo pr.
Patras, 7 June 1899, Heldreich (W-HAL) (ad P. gramineam trans.). Dhiakopton,
8 June 1899, Heldreich (W-HAL) (ad P. gramineam trans.). Between Dhiakopton
and Megaspelion, 19 Oct. 1939, Davis roo3 (E ; K) (ad P. gramineam trans.).
Panakhaikon Oros, Kokkinobryssi, 22 June 1949, Goulimy 50 (K).

TurKEY : Bosphore, Aucher-Eloy 553 (BM; K). Tokat, Niksar-Karakus,
5 Sept. 1954, Davis 24898 (E). Trabzon, Macka, to July 1934, Balls 1626 (E ;

154 REVISION OF THE GENUS PETRORHAGIA

K). Giresun, Tandere-Yavuzkenal, near Karinca, 13 Aug. 1952, Davis 20709
(BM; E; K). Rize, June 1866, René de Parquet (BM). Katahor, 7 July 1933,
Balls 474 (E ; K).

U.S.S.R.: Tiflis, Apr., Smirnoff (W-HAL) ; 29 May 1912, Holmberg 996 (W) ;
June 1881, Brotherus 153 (BM). Iberia, Aug. 1824, Wilhelms (K). Armenia,
Szovils 75 (K). Altaica, 1826, C. A. Meyer (BM).

IRAN : Twenty-five miles west of Zorab, 14 June 1929, Cowan & Darlington (K).
Mazanderan, Kudjur, inter Sanus et Kindj, 9-11 Aug. 1948, Rechinger fil. 6570
(BM ; K).

Introduced and naturalized :

SWEDEN : Skane, Paroecia Sirekopinge, 8 Aug. 1932, Samuelsson 769 (BM ;
W). Sireképing, Klasinge, 19 July 1901, Henningson (BM).

GREAT BriTAIN : Dorset : Abbotsbury, 5 Aug. 1925, Andrews (K). Pembroke :
Tenby, 2 Sept. 1909, Barley (K).

13b. Petrorhagia saxifraga var. glomerata (Ten.) Ball & Heywood, comb. nov.

Gypsophila rigida L., Sp. Pl. 1 : 408 (1753).

Gypsophila saxifraga var. glomerata Ten., Fl. Nap. 4 : 202 (1830).

Petrorhagia rigida (L.) Link, Handb. Erkenn. Gewachse 2 : 235 (1831).

Tunica rigida (L.) Reichb., Ic. Fl. Germ. & Helv. 6: 42 (1844).—Boiss., Fl. Or. 1: 518
(1867).

Very similar to var. saxifraga but flowers fasciculate.

IraLy : Venezia: Bassano, 15 Oct. 1854, Ball (E). Lazio : Roma, Collossea,
30 Aug. 1856, anon. (FI). Avellino, July 1888, Milansz (FI). Campania : Monte
Arasualo, 6 Oct. 1954, Agostini (FI). Inter Sorrento et Viso, 31 July 1874, Heldreich
(BM ; W-HAL). Puglia: Modugno (Bari), 25 Sept. 1938, Capztaneo (FI).
Calabria : Sila Fossiata (Longobucco), 29 July—3 Aug. 1918, Fort (FI).

Sicity : Palermo, June 1904, Ross 516 (E ; FI) ; 1893, Gasparini (BM ; CGE).
Etna, 1844, Tornabene (FI). Baida, 1848, Tineo (FI). Madonie, 1848, Tineo (FI).

JucosLtavia: Hrvatska: Rijeka (Fiume), 19 Oct. 1863, A. M. Smith (FI).
Makarska, Aug.—Oct. 1934, Gilliat-Smith 89 (K).

ALBANIA : Durrés, 30 Aug. 1935, Alston & Sandwith 2740 (BM ; K).

13c. Petrorhagia saxifraga var. gasparrinii (Guss.) Ball & Heywood, comb. nov.

Gypsophila gasparrinii Guss., Fl. Sicul. Synops. 1 : 474 (1843) (“ Gasparrini ”’).
Tunica gasparrinii (Guss.) Nyman, Consp. Fl. Eur. : 100 (1878).
Tunica saxifraga var. gasparrinii (Guss.) Fiori, Nuov. Fl. Anal. Ital. 1 : 505 (1924).

Stems procumbent with numerous rosettes of leaves. Lower leaves I-2:5 mm.
broad, oblong. Flowers solitary. Calyx 4-5-7 mm. long. Petals 7-10 mm. long.

Sicity : Messina, June 1866, Sequenza 81 (FI) ; Nicotra 21 (W-HAL). Isnello,
1848, Tineo (FI). Madonie, 1866, Pasquale (F1).

This very widespread species shows comparatively little variation throughout

REVISION OF THE GENUS PETRORHAGIA 155

its range. There is a certain amount of intergradation with P. graminea in
Peloponnisos and besides the typical variety there are the two varieties described
above, which occur mainly in Italy and Sicily.

Var. glomerata is characterized by the fasciculation of the flowers and this seems
to occur sporadically throughout the range of the species. However, in Italy
and Sicily it is much more frequent, and there tends to be a greater degree of fascicu-
lation. Gypsophila rigida L. appears to be identical with this variety. Var.
gasparrinu presents a different problem. This variety appears to be confined to
Sicily, and perhaps southern Italy. If it were not for this very restricted distribu-
tion var. gasparrinit would probably be regarded as no more than an abnormal
growth form, but in the present circumstances a field investigation seems to be
desirable to determine its true status. It is not at all clear whether var. glomerata
or var. gasparrinii occur in discrete populations at all, but what evidence is available
suggests that they may not.

Numerous other varieties have been described, but it seems doubtful whether
any of them deserve taxonomic recognition.

14. Petrorhagia graminea (Smith) Ball & Heywood, comb. nov. (Fig. 16.)

Gypsophila graminea Smith, Fl. Graec. Prodr. 1 : 279 (1809).
Tunica graminea (Smith) Boiss., Diagn. Pl. Or. Nov. 2 (8) : 60 (1849) ; Fl. Or. 1: 519
(1867).

Caespitose, usually very woody at the base with dense rosettes of leaves ; flower-
ing stems up to 40 cm. high, shortly and densely pubescent, sometimes glabrous
above. Basal leaves 8-20 mm. long, 0-5-0o-9 mm. broad. Inflorescence usually
lax, sometimes with a few flowers fastigiate ; epicalyx absent. Calyx 3°5-5:5 mm.
long, densely pubescent, sometimes with the teeth subglabrous ; costae 1-veined
with 2 obscure lateral veins ; teeth oblong, obtuse. Petals 5-10 mm. long, the
limb 1-8—-2:5 mm. broad, obcordate, pink, reddish, or white becoming pink. Seeds
I-I-1'5 mm. long, 0-8—0-9 mm. broad.

Distribution : Southern Greece (Peloponnisos).

GREECE : Peloponnisos : Messenia, Kalamata, Oct.—Dec. 1897, Heldretch 1410
(E; Kj; W-HAL); c. 1930, Guiol (BM). Elide, dans la forét de Danolada,
28 Oct. 1906, Maire & Petitmengin 2295 (W-HAL). Mt. Taygetos, Sept. 1929,
Guiol 1494 (BM).

This species, normally without epicalyx, appears to be very closely related to
P. saxifraga. A number of collections from the Peloponnisos are intermediate
between the two species, but it is not clear whether these represent occasional
hybrid swarms or geographically intermediate populations. Only an extensive
investigation in the area concerned is likely to provide a solution to this problem.

Bossier placed this species in his Tunica sect. Gypsophiloides, but the evidence
of intergradation with P. saxifraga quite clearly supports the transfer to Petrorhagia
sect. Petrorhagia. The seed characters, the white-hirtulose, oblong obtuse calyx
teeth, and the occasional fasciculation of the flowers also support this.

156 REVISION OF THE GENUS PETRORHAGIA

15. Petrorhagia rhiphaea (Pau & Font Quer) Ball & Heywood, comb. nov..

Tunica rhiphaea Pau & Font Quer in Font Quer, Iter Marocc. 1929: n. 147 (1930) ; in
Cavanillesia 3 : 77 (1930).

Stems 5-15 cm. high, glabrous in the lower part, glandular-pubescent at the apex.
Basal leaves 4-8 mm. long, 0-4—0-7 mm. broad, linear with obtuse apex. Inflorescence
lax ; epicalyx absent. Calyx 2:5-4:5 mm. long, glandular-pubescent ; costae
feebly 3-veined ; teeth ovate to oblong, obtuse. Petals 4-7 mm. long, the limb
0-8-1 mm. broad, linear-oblong, retuse, pale-yellow with purple veins. Seeds
(immature) 0-8-1-2 mm. long, 0-7—0-9 mm. broad.

Distribution : Spanish Morocco.

SPANISH Morocco : Inrupibus schistosis supra oppidum Mahzen dictum (Ktama),
ad 1,350 m., 29 June 1929, Font Quer 147 (BM, isotype).

This species is very similar to P. graminea and may eventually prove to be
identical with it, or perhaps a subspecies. It is kept distinct here owing to the
paucity of the material, and further collections are necessary before any definite
conclusions can be reached. The main differences are the smaller flowers with
pale-yellow, purple-veined (not pink, reddish or white becoming pink), much
narrower petals, and the shorter leaves.

Subsect. b. Thessalae Ball & Heywood

Petrorhagia subsect. Thessalae Ball & Heywood, subsect. nov.; a subsectione
Saxtfragis differt caulibus simplicibus vel sparse ramosis, floribus capitatis,
bracteis late ovatis vel suborbicularibus.

Type: P. thessala (Boiss.) Ball & Heywood.

Stems simple or with few branches. Flowers capitate ; bracts broadly ovate
or suborbicular, membranous and I-veined to brown-scarious and many-veined.

16. Petrorhagia thessala (Boiss.) Ball & Heywood, comb. nov. (Fig. 17.)

Tunica thessala Boiss., Diagn. Pl. Or. Nov. 2 (8) : 63 (1849) ; Fl. Or. 1: 518 (1867).

Gypsophila thessala (Boiss.) Nyman, Syll. Fl. Eur. : 238 (1854).

Tunica orphanidesiana Clem. in Mem. R. Accad. Sci. Torino, Ser. 2, 16 : 17 (1855) ; Sertul.
Or, 2 E7 (1855):

Stems 10-35 cm. high, simple or slightly branched above, shortly papillose-
pubescent at least at the base. Leaves up to 15 mm. long, linear. Heads up to
10-flowered ; outer bracts 6-many, the largest 6-10 mm. long, 2-5-4 mm. broad,
about equalling the flowers, ovate, brown-scarious with white-membranous margin,
rarely 5-7 mm. long, 1:5-2 mm. wide, ovate-lanceolate, and almost completely
membranous except for the vein. Calyx 5-5-7 mm. long, glabrous or papillose with
oblong, obtuse teeth. Petals 6-5-8 mm. long, linear-spathulate, entire, retuse or

rarely crenate, white with pink or purple veins. Seeds 1-8-2-3 mm. long, I-3-1-7 mm.
broad.

REVISION OF THE GENUS PETRORHAGIA 157

Distribution : Greece ; doubtfully in southern Jugoslavia.

GREECE : Séerea Ellas : Parnes, 24 Aug. 1911, Tunta 1279 (W-HAL) ; 28 June
1930, Guiol 1307 (BM) ; 17 July 1956, Rechinger fil. 18159 (W). Thessalia : Pelion,
supra urbem Volo, 28 July 1893, Leonis, in Haldcsy, Iter Graec. Sec. (K ; W-HAL)
Olympus, Aucher 537 (K, isotype). Olympi Thessaliae inter Hagios Elias et
Litochoren, July 1857, Orphanides (BM ; CGE ; E ; K ; W-HAL). Hagios Elias
prope Tyrnavo, 30 July 1885, Heldreich (W-HAL) ; 3 Aug. 1896, Sintenis 1106
(E; K; W-HAL). Milonna, 14 Aug. 1895, Formanek (W-HAL). Makedhonia :
In monte Boz-dagh prope Serrae, ad monast. Prodromos, 15 July 1936, Rechinger
fil. 10954 (W-RECH). Thraki: Near Oktchilar (Tochotai), Tedd 2041 (K).

This species has a somewhat disjunct distribution, its main centre being Olimbos
and neighbouring mountains, with apparently small populations on Parnis Oros, and
scattered localities in Macedonia and Thrace.

The plants from Tochotai (Oktchilar) differ somewhat from the rest of the species.
The bracts of the inflorescence are smaller and almost completely hyaline. They
approach P. macra and P. cyrenaica in this character, but otherwise resemble typical
P. thessala.

Records from Jugoslavia have not been confirmed, the specimens seen proving to
be Petrorhagia saxifraga. However, the distribution in Greece is such that the
species will probably be found to occur in both southern Jugoslavia and Bulgaria.

ae eS a
aes

[7 18

Fics. 17,18. 17, Petrorhagia thessala (Boiss.) Ball & Heywood. 18, P. dianthoides (Smith)
Ball & Heywood. a, calyx ; b, petal ; c, epicalyx bract or the largest bract from a head.

(All x5.)

158 REVISION OF THE GENUS PETRORHAGIA

17. Petrorhagia cyrenaica (Durand & Barratte) Ball & Heywood, sp. nov.

Tunica thessala var. cyrenaica Durand & Barratte, Fl. Lib. Prodr. : 33 (1910).—Maire, FI.
Afr, Nord10 > 281 (1963).

Stems 5-30 cm. high, glaucous, glabrous or papillose-scabrid at the base. Leaves
up to 15 mm. long, linear to setaceous. Heads 1~-3-flowered ; outer bracts
numerous, the largest 4-8 mm. long, 1:5-2:5 mm. broad, shorter than the flowers,
ovate-lanceolate, white-membranous with brown-scarious veins. Calyx 4-5-6 mm.
long, shortly and sparsely pubescent with oblong, obtuse teeth. Petals 5-5-8 mm.
long, linear-spathulate, ?entire, white, pale-red on the lower surface. Seeds
(immature) I-3-1-5 mm. long, 1:1 mm. broad.

Distribution : Libya.
LispyA. Zaouia Sidi-Mahdi entre Koubba et Lamloude, 13 May 1887, Taubert 559

(E, isotype). El Hania, 8 May 1934, Pampanini & Pichi-Sermolli 2577 (KK). Wadi
Derna, 8 Apr. 1939, Sandwith 2487 (K).

Very little material of this species has been seen, but it appears to be distinguish-
able from P. thessala by its fewer-flowered heads with smaller flowers, and smaller
almost entirely membranous bracts. Also, in the material seen, the calyx has
always been sparsely pubescent, although a glabrous variant has been described by
Pampanini (Arch. Bot. Forli 12: 24 (1936)).

18. Petrorhagia macra (Boiss. & Hausskn.) Ball & Heywood, comb. nov.
Tunica macra Boiss. & Hausskn. in Boiss., Fl. Or., Suppl. : 81 (1888).
Tunica gracilis F. N. Williams in Journ. of Bot. 28 : 196 (1890).

Stems 25-40 cm. high, slightly branched above, pubescent at the base, glabrous
elsewhere. Leaves up to 10 mm. long, oblong-linear to subulate. Heads 1-5-
flowered ; outer bracts 4-many, the largest 3-5-5 mm. long, 0-7-1:3 mm. broad,
triangular-lanceolate to ovate, acuminate, white-membranous with a brown-scarious
vein. Calyx 5-5-8 mm. long, sparsely pubescent with more or less oblong-obtuse
teeth. Petals 8-9 mm. long, linear-spathulate, entire, white. Seeds not known.

Distribution : Iran (Kurdistan).

IRAN: Monte Shahu, 1867, Haussknecht (K, isotype, also holotype of Tunica
gracilis).

Very similar to P. thessala and P. cyrenaica. Tunica gracilis was described from
isotype material of T. macra. The specimen closely fits the description of T. macra,

and there does not seem to be any reason why it should be regarded as a distinct
species.

19. Petrorhagia dianthoides (Smith) Ball & Heywood, comb. nov. (Fig. 18.)

Gypsophila dianthoides Smith, Fl. Graec. Prodr. 1 : 280 (1809).

Tunica dianthoides (Smith) Fisch. & Meyer, Index Sem. Hort. Bot. Petrop. 4 : 48 (1837).—
Boiss., Fl. Or. r : 518 (1867).

Kohlrauschia sibthorpit Kunth, Fl. Berol. Fam. Nat. Dispos. 1 : 109 (1838), nom. illegit.

Stems up to 40 cm. high, simple or slightly branched above, glabrous. Leaves up

REVISION OF THE GENUS PETRORHAGIA 159

to 20 mm. long, linear to subulate. Heads 1-6(-8)-flowered ; outer bracts 4—many,
the largest 4:5-10 mm. long, I-2:5 mm. broad, lanceolate to ovate-lanceolate, white-
membranous except for the brown-scarious vein. Calyx 5~7 mm. long, pubescent,
with triangular-lanceolate, more or less acute teeth. Petals 6-10 mm. long, oblong-
spathulate, entire, white with red veins. Seeds (immature) I-1-3 mm. long,
o-8—o-9 mm. broad.

Distribution : Crete.

CRETE : Mont. Sphacioticorum, July 1846, Heldreich (BM; CGE; E; K;
W-HAL ; isotypes). Hag. Roumeli et Samaria, dist. Sphakia, 12-13 July 1899,
Baldacci 21m (BM ; W-HAL) ; June 1932, Guwiol 2125 (BM). Aradhena, 24 July
1882, Spreitzenhofer (W-HAL) ; July 1932, Atchley 1694 (K). Paleocastis, Sitia,
May 1852, Heldreich (E). Theriso gorge near Cnea, 22 July 1938, Davis 303 (BM ;
E ; K). Imbros gorge, 25 May 1938, Davis 185 (E).

This species appears to be one of the most distinct in the section,

Sect. 4. DIANTHELLA (Boiss.) Ball & Heywood
Petrorhagia sect. Dianthella (Boiss.) Ball & Heywood, comb. nov.
Tunica sect. Dianthella Boiss., Fl. Or. 1 : 517 (1867).

Type : Tunica pamphylica Boiss. & Balansa (= P. pamphylica (Boiss. & Balansa)
Ball & Heywood).

Annual. Leaves 3-veined. Flowers solitary or subcapitate ; bracts ovate,
brown- or purplish-scarious, 3-many-veined. Calyx costae 3-7-veined. Petals not
clawed. Seeds small, blackish-brown, reticulate-tuberculate, with thickened margin.

20. Petrorhagia pamphylica (Boiss. & Balansa) Ball & Heywood, comb. nov.
(Fig. 19.)
Tunica pamphylica Boiss. & Balansa in Boiss., Diagn. Pl. Or. Nov. 3 (6) : 27 (1859).—
Boiss, Fly Ort = 517-(1867),

Stems 10-35 cm. high, pseudodichotomously branched from the base, glabrous.
Cauline leaves up to 16 mm. long, I mm. broad, linear, 3-5-veined. Flowers
solitary ; epicalyx bracts 6-12, 3-4 mm. long, shorter than the calyx, oblong to
ovate, acute, brown- or purplish-scarious with white-membranous margin and with
5-many thick veins. Calyx 5-7 mm. long ; costae very broad, 6~7-veined, the
commisural membranes almost absent ; teeth o-8-1-1 mm. broad, lanceolate-
acuminate. Petals 7-9 mm. long, the limb c. 0-5 mm. broad, linear-spathulate,
obtuse, entire, pale-pink. Seeds 1-6-2 mm. long, 1-2-1-7 mm. broad.

Distribution : Turkey (southern Anatolia).
Turkey : Antalya, near Gebiz, stony hillside, 1,000 m., 22 July 1949, Davis 15478
(BM s Bue).
21. Petrorhagia peroninii (Boiss.) Ball & Heywood, comb. nov. (Fig. 20 ;
Plate 14 C.)
Tunica peroninii Boiss., Fl. Or., Suppl. : 81 (1888) (“ Peronini’’).

160 REVISION OF THE GENUS PETRORHAGIA

Stems 15-30 cm. high, pseudodichotomously branched from the base, pruinose-
papillose. Cauline leaves 8-20 mm. long, c. 1 mm. broad, linear, 3-veined. Flowers
solitary or subcapitate ; bracts and epicalyx bracts 6—-many, up to 5-7-5 mm. long,
exceeding the calyx, lanceolate, brown- or purplish-scarious with white-membranous
margin, papillose, 3(—5)-veined. Calyx 5-7 mm. long ; costae 3-veined, with a
narrow but distinct commissural membrane ; teeth 0-4-0-7 mm. broad, oblong-
lanceolate. Petals 5:5-8-5 mm. long, the limb c. 0-5 mm. broad, linear-spathulate,
obtuse, entire, white, sometimes purplish on the lower surface. Seeds I-I-1-:2 mm.

long, 0-7-0-9 mm. broad.

--
es Fania)

19 20

19, Petrorhagia pamphylica (Boiss. & Balansa) Ball & Heywood. 20,

FIGs. I9, 20.
a, calyx ; b, petal ; c, epicalyx bract. (All x5.)

P. peroninii (Boiss.) Ball & Heywood.

Distribution : Turkey (southern Anatolia).

TuRKEY : Prov. Mersin, Dist. Anamur, montagne ouest d’Anamour, June 1872,
Peronin 5 (BM; K> isotypes). Between Ferhenk and Anamur, rocky meta-
morphic hills, 50 m., 19 Aug. 1949, Davis 16329 (BM; E; K).

These two rare species, as already mentioned (p.126), appear in some respects to
form a link between Petrorhagia and Dianthus.

Sect. 5. KOonLRAUSCHIA (Kunth) Ball & Heywood

Petrorhagia sect. Kohlrauschia (Kunth) Ball & Heywood, comb. nov.

Kohlrauschia Kunth, Fl. Berol. Fam. Nat. Dispos. 1 : 108 (1838).

Dianthus sect. Kohlrauschia (Kunth) Fenzl in Endl., Gen. Pl. : 971 (1840).
Tunica sect. Kohlrauschia (Kunth) Griseb., Spicil. Fl. Rumel. 1 : 185 (1843).
Dianthus subgen. Proliferastrum F. N. Williams in Journ. of Bot. 23 : 347 (1885).

Type : Kohlrauschia prolifera (L.) Kunth (= P. prolifera (L.) Ball & Heywood).

Annual. Leaves 3-veined. Flowers capitate ; bracts very broad, orbicular-
ovate or suborbicular, brown-scarious, many-veined. Petals distinctly clawed.
Seeds small, blackish-brown, reticulate to papillose, with thickened margin.

REVISION OF THE GENUS PETRORHAGIA 161

22. Petrorhagia prolifera (L.) Ball & Heywood, comb. nov. (Plate 15 A.)
Dianthus prolifer L., Sp. Pl. 1: 410 (1753).
Dianthus diminutus L., op. cit., ed. 2, I : 587 (1762).
Tunica prolifera (L.) Scop., Fl. Carniol., ed. 2, 1 : 299 (1772).
Caryophyllus aridus Moench, Meth. Pl.: 59 (1794), nom. illegit.
Kohlrauschia prolifera (L.) Kunth, Fl. Berol. Fam. Nat. Dispos. 1 : 10g (1838).
Kohlrauschia diminuta (L.) Reichb., Ic. Fl. Germ. & Helv. 6: 43 (1844).

Stems up to 50 cm. high, usually simple, glabrous or scabrid. Cauline leaves up to
40 mm. long and 2 mm. broad, linear-oblong, 3-veined ; sheath 1-2-5 mm. long,
about as long as broad. Largest bracts of head 6-12 mm. long, 3~7(—8) mm. broad,
usually obtuse. Calyx 10-13 mm. long ; costae 3—veined ; teeth oblong, obtuse.
Petals 10-14 mm. long, the limb 2-3-5 mm. wide, obcordate, pink or purplish. Seeds
I-3-I'9 mm. long, 0-8—1-I mm. broad, reticulate.

Distribution : Central Europe, mountains of southern Europe, Caucasus, Turkey
(northern Anatolia), mountains of western North Africa.

SWEDEN : Paroecia Garde, Petard, Gotland, 30 June 1926, Samuelsson 797 (BM ;
W). Prope Resmo, Oland, 15 July 1860, July 1864, Ahlberg (BM ; CGE; K).

DEnMARK : Ins. Samsoe, 28 July 1887, Jensen (W-HAL). Ins. Bornholm, July
1848, Lange (CGE). Rorvig, Aug. 1866, Steenhiip (E).

GERMANY : Sandhausen, 8 km. S. of Heidelberg Baden, 1 Aug. 1935, Stearn 70
(K). Bei Mannheim, 2 Aug. 1859, Schweinfurth (BM). Frankfurt, 24 June 1856,
Bagge (BM). Bonn, 16 June 1851, Blackie (E). Schleswig-Holstein, June-July,
Hansen 347 (BM). Calshalle, Kreuznach, 2 Sept. 1868, Tvimen (BM). S. von
Aschaffenburg (Bavaria), July 1905, Rubner (E). Prope K1.-Kreutz, Brandenburg,
Aug. 1922, Gort (K).

PoLanp : Krappilk (Schlesien), 19 July 1899, Ziesché (E).

CZECHOSLOVAKIA : Litomefice, 17 Sept. 1918, Korb (W). Pressburg, 7 Sept.
1913, Korb (W). Westfusse der Kahlberges (B.-Leipa), Aug. 1910, Petrak 330
(BM ; E). Prope Veveri Bytyska, Moravia occid., Aug. 1924, Sivjaev 26 (K).

AustrRiA : Tirol: Boyen, Grabmayr (K). Nieder-Osterreich : Wiener-Neustadt,
Kerner, Fl. Austro-Hung. 2509 (BM; K; W-HAL; W-RECH). Prope Wien,
Aug. 1878, Halécsy (W-HAL). Baden, 18 July 1900, Krebs (E). Burgenland :
Leithagebirge, Naglersberge, 21 June 1923, Korb (W). Am Eisenbahndamm
zwischen Weiden und Neusiedler See, 21 Sept. 1924, Rechinger (W-RECH).

Huncary : Ins. Csepel, 2 July 1913, Filarszky & Kiimmerle, Fl. Hung. Cent. IT, 90
(BM; E; K; W-RECH). Pest, June 1872, Richter (BM). Bei Magyarfalva,
24 June 1905, Vetter (W).

RoMANIA : Baile Herculane, Caras Severin, 13 June 1922, Bujorean, Fl. Rom. 4or
(BM ; E; K). Carmen Sylva, Constanta, 14 Aug. 1931, Enculescu, Fl. Rom. 4o1b
(K).

BULGARIA : Ladovo, 8 June 1898, Sivibrny (E). Dragoman, mont. Golem Cepan,
31 July 1930, Rechinger fil. (W-RECH). Djenden Tepe, bei Philipopel, 25 July

162 REVISION OF THE GENUS PETRORHAGIA

1891, Gheorghieff (W-HAL). Dolni Pasarel, 6 July 1957, Newman (K). Prope
Varna, 16 July 1930, Rechinger fil. 1021 (W-RECH). Bei Tirnorvo, 1896, Urumoff
8r (W-HAL). Bei Lowtscha, 1895, Urumoff (W-HAL). Supra urbem Kolarov-
grad, 9 July 1951, Zelezova, Pl. Bulgaricae Exs. 25 (K). Gorne Banja, 20 June 1921,
Uromov (K). Plovdiv, 20 July 1930, Rechinger fil. 1165 (W-RECH). Near R.
Topolnitza, 36 km. from Pazardzhik, 16 July 1950, Mason 27 (K). Prope Lakatnik,
Stara Planina, 29 July 1930, Rechinger fil. 1760 (W-RECH). Deli Orman prope
Russe (Rustschuk), 13 July 1930, Rechinger fil. 747 (W-RECH).

JUGOSLAVIA : Makedonija: Mte. Ohrida, 1908, Dimonie (W-HAL). Veles,
Ig June 1905, Adamovié (E). Bei RaduSe, 11 July 1937, Behy (W-RECH). Scardus
geb., July 1876, Hertz (K). Gjefgjeli, 1-2 June 1927, Rechinger fil. 1577
(W-RECH). Bosna 1 Hercegovina: M. Bokievria prope Astrozai, 4 July 1926,
Maly (K). Moscanica, 4 July 1933, Guilliat-Smith 3266 (K). Mostar, 8 July
1955, Hepper 870 (K). Srbija: Prope Vranja, 18 June 1900, Adamovié (E).
S. of Streocha (Streoci), Prokletije, 18 July 1956, Carpenter 340 (K). Visoki Decani,
prope Peé, 17-19 July 1933, Rechinger fil. & Scheffer 1070 (W-RECH). Fluv.
Pecska Bistrica prope Pe¢, 3 July 1933, Rechinger fil. & Scheffer 121 (W-RECH).
Dalmacya : Makarska, 1934, Jozipovié (K). Ragusam (Dubrovnik), June 1907,
Adamovié (K). Napoda, Kriodsijc, 6 June 1906, Schneider (W). Popovici (Tepui
Dol), 20 July 1926, Lorchuigg (W-RECH). Hrvatska : Vernde, Istria, 17 May 1875,
Freyn (K).

ALBANIA : Sarandé, 10 July 1932, Alston & Sandwith (BM). Mali Dajtit near

Tirana, 9 Aug. 1938, Hepburn 26 (K). Near Richa, 4 July, Giuseppi (K). Selce,
dist. Klementi, 2 Aug. 1901, Baldacci 192 (BM).

GREECE : Sterea Ellas : Mavrolitharion to Mousonitsa (Oiti), 14 June 1937, Balls
& Gourlay 3292 (BM; K). Ad Paliampela (Akarnania), 22 July 1889, Baldacci
(K ; W-HAL). Ifivos: Janina, Sept.—Oct. 1895, Baldacci (BM); July 1931,
Guiol (BM). Prope Dramissous, June-July 1929, Guiol (BM). Mt. Peristeri prope
Syraka, 14 July 1893, Haldcsy (W-HAL) ; Aug. 1931, Atchley 992 (K). M. Murga
(Tzoumerka), 6 July 1895, Baldacci 15 (K). Montes Pindus, prope Metsovon,
28 July 1956, Rechinger fil. 18298 (LIVU). Makedhonia: Lacus Dojran, 6 June 1936,
Rechinger fil. 9147 (W-RECH). Boz-dagh prope Serrai, 15 July 1936, Rechinger fil.
11050 (W-RECH). Samarina to Perevali, 14 July 1937, Balls & Gourlay 3609 (E ;
K). Near Thessaloniki, 19 Sept. 1917, Ramsbottom (BM). _ S. slopes of Krousia
Oros, 30 June 1917, Turrill (BM). Thraki: Fluv. Nestos (Mesta) prope Toxotai,
12 June 1936, Rechinger fil. 9279 (W-RECH). Prope Alexandroupolis, 29 May
1934, Rechinger 6054 (W-RECH). Xanthi, 23 June 1931, Tedd 666 (K). Derecol
valley near Xanthi, 4 Oct. 1929, Tedd 70 (K).

SWITZERLAND : Between Sierre and Vissoie, Valais, 3 Aug. 1903, Lester-Garland
(K). Prope Geneva, Boissier (K). Martigny, 27 Aug. 1858, Fraser (E). Lausanne,
June 1879, Favrat & Barbey (K). Argau, Unteres Aaretal, 30 June 1954, Koch
(W). Cressier, 10 June 1834, Neuchatch (E).

ItaLy : Basilicata: Pignola, 8 June 1924, Gaviolo (FI). Gruppo del Pollino,

REVISION OF THE GENUS PETRORHAGIA 163

19 July 1935, Gavioli (FI). Luconia, M. San Michele, 2 July 1932, Gavioli (FI).
Campania : Monte Alburno, Da Corleto M. a C. Sierre, 12 Aug. 1936, Philippis (FI).
Presso Caserta, June 1870, Tenace (FI). Lazio : Tivoli, Sul Monte Catillo, 12 June
1904, Vaccari (FI). Monte Albani, Nemi, 20 June 1875, Marchesetit (FI). Montero-
tondo (Roma), 22 Oct. 1950, Steinberg (FI). Umbria : Stroncone, 22 Sept. 1893,
Mari (FI). Pemgin, 30 Oct. 1947, Bettini (FI). Venezia: Mont. Ricco prope
Monselice, May 1911, Béguinot 1657 (BM ; FI; K). Verona, 27 July 1888, Caruel
(FI). Romagnano, Aug. 1888, Goivan (FI). Tirolo Italiano: S. Leonardo in
Passiria, Alto Adige, 15 Aug. 1949, Zenari (FI). Cembra, 20 Sept. 1948, Whitehead
(K). Emilia: M. Titano, San Marino, 25 June 1912 and 15 Oct. 1916, Pampanini
(FI). Osservanza, presso Bologna, June 1900, Ceroni (FI). Spezzano, 19 July 1882,
Fiori (FI). M. Chiarugola, Parma, 18 Aug. 1919, Mimo (FI). Marche: Mt.
Fortino, Apennini Picenim, 1836, Marzialetti (FI). Mt. Catria, 23 July 1866,
Parlatore (FI). Nevola e Mifr, 4 July 1934, Bettina (FI). Puglia : prope Gallipoli,
June 1883, Groves (FI). Abbruzzo : Monte Morrone, 1891, Profeta (FI). Majella,
30 June 1905, Vaccari (FI). Prope Gagliano, Sirente, July 1876, Groves (FI),
Lombardia: Malgrate, 1 Oct. 1893, Camperio (FI). Grumone, 3 Sept. 1864.
Parlatore (¥F1).

SicILy : Sagana, 12 Aug. 1836, Parlatore (FI). Etnae, Sept. 1844, Tornabene
(FI). Maudenici, 4 June 1882, Borzi (FI).

Corsica: Bastia, 28 Sept. 1880, Chubern (FI). Bastelica, 20 June 1878,
Reverchon (K).

FRANCE : Muerthe-et-Moselle : Nancy, 1871, Gamble (K). Jura: Serre, Sept.
1866, Blanche. Savoie: Prés Albens, 28 Sept. 1881, anon. (K). Hautes-Alpes : Gap,
3 July 1871, Reverchon (K). Briancon, 18 July 1871, Reverchon (K). Embruin, Sept.
1887, Gamble 19660 (K). Iséve : Pont de Beauvois, Sept. 1860, anon. (K). Grenoble,
June 1840, Haden (BM). Puy de Dime: Puy de Montaudon, 16 Aug. 1927, Burtt-Davy
167/27 (K). Rhéne: Lyon, 1852, Jordan (K). Céte d’Or: Dijon, 1 Aug. 1894,
Clarke 47629 (K). Somme: St. Valery-sur-Somme, 10 July 1959, Carter (K).
Niévre : Nevers, Aug. 1840, J. Ball (E). Eure: Andelys, Aug. 1844, Mill (BM ; K).
Orne: Alencon, 28 June 1886, Beaudoin (LIVU). Loire-Inférieure : Nantes, 1897,
Blow (E). Seine-et-Oise: Point du Jour, prés Paris, 4 July 1875, Bonnet (K).
Seine-et-Marne : Fontainbleau, Henslow (E). Haute-Garonne : Toulouse, 25 July
1952, Billot 2027 (BM ; LIVU). Hérault : W. of Montpellier, 14 Sept. 1953, Rawden
48 (K). Pyrénées-Orientales : Verneb, 15 July 1934, Wyatt (K). Hautes-Pyrénées :
Gedre, July 1870, Bordére (K). Basses-Pyrénées : Laruns, 28 July 1922, Hutchinson,
Matthews & Riley 290 (K).

BELGIUM : Mariembourg, 23 June 1935, Mosserony (K).

LuxemBourc : N. du Hobscheid, 18 June 1953, Reichling (BM). Eich, 23 Sept.
1925, Rundberg (W-RECH).

Spain : Ur, Cerdagne, 21 Aug. 1956, Hooper (K). San Mateo (Castellén de la
Plana), 1954, Rechinger fil. (W).

164 REVISION OF THE GENUS PETRORHAGIA

Morocco: Fimelil, Atlas, 12 June 1926, Lindberg 3880 (K). Ras-el-Ma, prope
Axron, 1926, Lindberg 4293 (W).

U.S.S.R. : Crimea, 1855, Munro (K). Bei Sudak, 13 July 1896, Callier 378 (E ;
W-HAL ; W-RECH). Lagodechi, fluv. Lagodech, Caucasus Kachetia, 17 July
1898, Tomin (E).

TurKEY : Adana, Feke, Sencan Deresi between Giiriimze and Siiphandere,
I July 1952, Davis 19603 (K). Adapazari, Arifiye, 1 July 1962, Davis 36281 (E).

Introduced :

GREAT BRITAIN : Sussex : Lewes, 1902, Davy (BM). Norfolk : near Northwold,
19 Sept. 1927, Little (BM).

The typification and relationships of this species are discussed by Ball & Heywood
(1962).

23. Petrorhagia nanteuilii (Burnat) Ball & Heywood, comb. nov. (Fig. 21 ;
Plate 15 B.)
Dianthus nanteuilii Burnat, Fl. Alp. Marit. 1 : 221 (1892).
Tunica nanteuilii (Burnat) Giirke in K. Richt., Pl. Eur. 2 : 338 (1903).
Tunica prolifera var. nanteuilit (Burnat) Briq., Prodr. Fl. Corse 1 : 569 (1910).
Tunica prolifera subsp. nanteutlii (Burnat) Graebn. in Aschers. & Graebn., Synops. Mitteleur.
El. 5 :(2) = :264: (t921):
Kohlrauschia nanteuilii (Burnat) Ball & Heywood in Watsonia 5 : 115 (1962).

Very similar to P. prolifera but the stems sometimes pubescent in the middle ;
leaf sheaths 1-3-5 mm. long, sometimes up to twice as long as broad ; seeds
tuberculate.

Distribution : Western Europe and western North Africa.

GREAT BRITAIN: Isle of Wight: Douvre Common, Aug. 1841, C. A. Wright
(BM). Ryde, 6 July 1852, More (BM). Hampshire : Hayling Island, 18 Aug. 1922,
Foggitt (BM). Portsea Island, 1890, Snope (LIVU). Sussex : Selsey, 1837, Forster
(BM). E. of Pagham, 21 June 1914, Salmon (BM). Pagham, 25 June and 29 July
1901, Marshall (LIVU). Kent: Dungeness, July 1903, Chandler (BM). Hythe,
Aug. 1903, Riddelsdell (BM). Norfolk: Stoke Ferry, July 1890, Aug. 1891, Cross
(BM).

CHANNEL ISLANDS: Jersey: Quenvais, 14 July 1901, Lester-Garland (LIVU).
St. Ouens Bay, 14 June 1862, Briggs (LIVU). St. Brelade’s Bay, 8 June 1884,
Hanbury (LIVU). Guernsey : C. 1858, Brewer (BM).

FRANCE : Iséve : South of St. Marcellin, 12 Sept. 1953, Rawdon 20 (K). Coétes
du Nord: St. Lunaire, 1883, Lester-Garland (K). Loire Inférieure : Pointe de
Chemoulin, 1871, Genevier (BM ; CGE). Lot: Prés de Cahors, June 1855, Bullot
2027b (BM ; CGE ; LIVU). Guronde : Bordeaux, 1896-97, Neyraut (K ; W-HAL).
Hautes-Pyrénées : Luz St. Sauveur, 18 Aug. 1932, Meinertzhagen (BM). Prope
Cauterets, July 1845, Spruce (CGE ; K).

Corsica : Prés Bastia, 17 June 1881, Chubern (FI).

SARDINIA : Isola Maddalena, June 1893, Vaccari (FI).

REVISION OF THE GENUS PETRORHAGIA 165

SPAIN : Versus Vallvidrena, prope Barcelone, June 1876, Tvemols (W-HAL).
S. Juan de Alcaraz, 16 June 1860, Bourgeau 963 (CGE ; E; K). Bayona, Ponte-
vedra, 20 June 1935, Schrafer (BM). Madrid, June 1919, Sennen 3662 (K). S.
Guadarrama, 5 June 1933, Afchley 123 (K). Above Cercedilla, S. Guadarrama,
7 June 1926, Wilmott (BM). Alcantara Bridge, Toledo, 6 June 1926, Wilmott (BM).
Despefiaperros, 2 June 1927, Wilmott & Lacaita (BM). Venta de Cardenas, 11 June
1952, Rotvainen (K). Slopes of Yelmo Grande, Sierra de Segura, 22 June 1955,
Heywood 2802 (LIVU). El Campillo, Sierra de Segura, 21 June 1955, Heywood 2670
(LIVU). Barranco de la Cueva de los Cervales, S. Magina, 6 June 1925, Cuatrecasas
5311 (K). Minas de Beires, S. Nevada, 20 June 1926, Wilmott & Lofthouse (BM).
Supra Guejar, S. Nevada, 22 Aug. 1844, Willkomm 136 (K). Prados del Aire,
S. Nevada, 10 June 1926, Wilmott & Lofthouse (BM). Ronda, 7 May 1924, Ellman
& Hubbard 231 (K). Gibraltar, 5 June 1913, Wolley-Dod 2113 (K). Guadacorte,
17 May 1912, Wolley-Dod 1060 (BM). Prope San Roque, 16 June 1871, J. Ball (K).
Almendral (Cadiz), 18 May 1957, Taylor (K).

2| 22

Fics. 21, 22. 21, Petrorhagia nanteuilii (Burnat) Ball & Heywood. 22, P. glumacea
(Bory & Chaub.) Ball & Heywood var. glumacea. a, calyx ; b, petal ; c, epicalyx bract
or the largest bract from a head. (All x5.)

PorTUGAL : Coimbra, 5 July 1887, Murray (BM). Estacao Velha, Coimbra,
3 June 1952, Matos (BM). Peneda de Saudade, Coimbra, June 1886, Moller
(W-HAL). Near Lisbon, 1935, Scarlett 180 (K). Moita, June 1840, Welwitsch 222
(CGE ; K). Tapada da Ajuda, 5 June 1933, Atchley 256 (K) ; 3 May 1940, P. Silva

166 REVISION OF THE GENUS PETRORHAGIA

(K). Villa Nova de Rainha, June 1846, Welwitsch (BM; K). Near Oporto,
16 July 1887, Murray (BM). Caldar de Gerez, June 1888, Tait (BM).

Morocco: Tichka l’Agadal, 10 June 1936, Balls 2766 (BM; K). Amizmiz,
17 June 1936, Balls 2851 (BM ; K).

MADEIRA: Madeira, Lindley (K) ; July 1862, Clarke (K). Funchal, 27 Mar.
1949, Sledge (BM). Bay E. of Pont Gorda, 27 Apr. 1924, Riley (K).

CANARY IsLANDS: Tenerife: Teneriffae, Webb (K). Cuevas Negras, 27 Apr.
1845, Bourgeau 709 (K). Santa Ursula, 5 Mar. 1933, Asplund rog (K). Montana
de la Horca Orotava, 26 May 1913, Sprague & Hutchinson 4o (K). Barranco del
Carmen, 31 May and 1 June 1913, Sprague & Hutchinson 163 (K), 170 (K). La
Palma: El Paso, 10 June 1913, Sprague & Hutchinson 303 (K).

For full discussion of the separation of this species from P. prolifera see Ball
& Heywood (1962).

24. Petrorhagia velutina (Guss.) Ball & Heywood, comb. nov. (Plate 15 Cc.)

Dianthus velutinus Guss., Index Sem. Hort. Bocc. 1825 : 2 (1825).

Tunica velutina (Guss.) Fisch. & Meyer, Index Sem. Hort. Petrop. 6 : 66 (1839).
Gypsophila velutina (Guss.) D. Dietr., Synops. Pl. 2: 1542 (1840).

Kohlrauschia velutina (Guss.) Reichb., Ic. Fl. Germ. & Helv. 4: 43 (1844).

Stems up to 50 cm. high, usually simple, usually with crowded glandular hairs on
the middle internodes, rarely almost glabrous. Cauline leaves up to 35 mm. long and
2mm. broad, linear-oblong, 3-veined ; sheath 3-7(-8) mm. long, at least twice as
long as broad. Largest bracts of head 7-12 mm. long, 4:5-8-5 mm. broad,
mucronate. Calyx 8-14 mm. long ; costae 3-veined ; teeth oblong, obtuse. Petals
11-16 mm. long, the limb 1-2-2-5 mm. broad, obcordate, sometimes bifid, pink or
purplish. Seeds 1-1-3 mm. long, 0-7-0-8 mm. broad, strongly tuberculate or
papillose.

Distribution : Mediterranean region.

PorTuGAL : Alemtejo, Redondo, May 1891, Moller 1086 (W-HAL). Prope Villa
Nova, 1848-50, Welwitsch 1158 (BM).

SPAIN : Vega de Granada, June 1864, Campo (BM). Sierra de Chiva, May 1844,
Willkomm (BM). Barranco de Ohanes, Sierra Nevada, 21 June 1926, Wilmott &
Lofthouse (BM). Near Venta de Saladillo (Gibraltar), 15 Apr. 1912, Wolley-Dod 651
(BM). Almendral (Cadiz), 18 May 1957, Taylor (K).

FRANCE : Alpes-Maritimes: Prés Nice, 7 May 1861, Bourgeau (FI). Esteril,
7 May 1889, Bonafous & Vidal (K).

Corsica : Pentes du Pigno, a Bastia, 20 June 1865, Mabille, Herb. Cors. 73
(BM ; CGE; FI; K). Bastelica, 20 June 1878, Reverchon (BM ; K ; W-HAL). Is.
Sanguinare, 25 May 1848, Reguien (K). Ajaccio, Apr. 1899, Collett (K). Near
Corte, Porter 176 (BM). Near Porto, 22 Apr. 1928, Edwards (BM). Travo, 29 Apr.
1930, Bates (BM).

Iraty: Liguria: Pizzo, Apr.-May, Ricca (FI). Emilia: Bologna, Monti

REVISION OF THE GENUS PETRORHAGIA 167

Sabbione, 11 June 1873, Marchesetti (FI). Presso Modena, 27 June 1808, Fiori (FI).
Marche: Flora Picena, Orsini (FI). Toscana: Grosseto, 9 Apr. 1843, Parlatore
(FI). Isola del Giglio, 24 Apr. 1895, Sommer (FI). Careggi, 13 May 1866, Bucci
(FI). Pisa, 15 May 1870, Groves (FI; K) ; May 1856, Savi in Billot 2028 (BM).
Firenze, 29 Apr. 1874, Leviery (CGE ; FI; K; W-HAL). S. Vincenzo, 2 May
1868, Parlatore (FI). Presso Buggiane, 1 Apr. 1933, Picht-Sermolli (FI). Colli di
Terontola (Arezzo), 20 May 1916, Fiori (FI). Lazio: Lago d’Agnano, 1g Apr.
1841, Heldreicch (CGE; FI; K). Civita Castellana, 11 May 1847, Bentham (K).
Roma, 17 May 1903, Vaccari (FI). Campania: Licola, 8 Apr. 1955, Agostini (FI).
Monte Alburno, 29 May 1951, Philippis & Moggi (FI). Napoli, May 1889, Guadagno
(FI). Mts. di Agerola, 15 May 1911, Guadagno (FI ; W-HAL). Between Amalfi
and Salerno, June 1883, Lacaita (K). Lipari, Apr. 1902, Zodda (FI). Insula
Caprearum, 8 May 1878, Heldreich (W-HAL). Puglia: Monte Calvo (Gargano),
27 May 1893, Martelli (FI). Mater Grazia, Leucarpide, Apr. 1877, Profeta (FI).
Basilicata : Potenza, 13 June 1926, Gavioli (FI). Calabria : Stilo et Mongiana,
18 June 1877, Biondi (FI). Prope Pezzo, 3 May 1877, Biondi (FI). Rossano, 4 May
1872, Aint: (FI). Palene, 20 May 1877, Arcangeli (FI). Serra San Bruno, 30 Apr.
1884, Zwierlein (FI).

SARDINIA : Capo Carlo, 12 Apr. 1894, Martelli (FI). Mte. Genargentu, 21-22 June
1897, Martelli (FI). Isola dei Cavoli, 12 Apr. 1894, Martelli (FI). Isola Maddalena,
Apr. 1893, Vaccari (FI). Tempio, 23 May 1881, 3 June 1882, Reverchon 68 (E ;
FI; K; W-HAL). Orune, May 1899, Martells (FI). Sassari, 7 May 1895,
Martelli (FI). Cagliare, Mar. 1828, Thomas (K).

SiciLy : Madonie, Castelvetrano, 1842, Parlatore (FI). Palermo (Panormitani),
May, Todaro 437 (BM; CGE; FI; K; W-HAL). Chiani di la Penna, June,
Mina 330 (FI). Piana del Greci, 21 May 1855, Huet du Pavillon (BM ; CGE ;
FI; K). Messina, Canepo, 1865, Seguenza (FI). Pantelleria, Apr. 1890, Ross (FI).
Castanea, May 1867, Mallandrino (FI).

JUGOSLAVIA : Hrvatska: Pola (Pula), 7 June 1874, Freyn (FI). Promontore
(Istria), 23 May 1898, Marchesetti (FI). Fasana, 10 June 1909, Korb (W).

ALBANIA : Sarande, 3 June 1933, Alston & Sandwith 1372 (BM ; K). Levani,
23 Apr. 1918, Schneider (W). Ljusme, 1 May 1918, Schneider (W).

Burearia : S. of Varna, 20 May 1925, Gilliat-Smith 1134 (K).

GREECE : Thraki: Dedeagach (Alexandroupolis), 7 May 1891, Sintenis &
Bornmiiller 75 (K ; W-HAL). Porto Lago, 17 Apr. 1933, Tedd 939 (K). Near
Xanthi, 21 May 1929, Tedd 8 (K). Makedhonia : Cassandra, 4 May 1891, Abd-ur-
Rahman Nadji (W-HAL). Simopetra, Athos, 13 Apr. 1934, Hill, Sandwith &
Turrill 2337 (K). Karadagh inter Thessaloniki et Serrai, 8 June 1936, Rechinger fil.
9214 (W-RECH). Near Snevce, Apr.-May 1918, Cook (BM). Between Kalamaria
and Mikra Bay, Mar. 1918, Ramsbottom (BM). Dora Tepe, E.N.E. of Doiran, 1918,
Gooding (BM). Antartikon, 30 May 1932, Alston G& Sandwith 59 (BM; K).
Vermion prope Naoussa, 30 May-1 June 1936, Rechinger fil. 8757 (W-RECH).
Thessalia: Kalampaka, 20 May 1896, Sintenis 251 (W-HAL). Sterea Ellas :

168 REVISION OF THE GENUS PETRORHAGIA

Korax prope Musinitza, 5 June 1899, Dérfler 283 (W-HAL). Agrapha, circa
monasterium Korona, 20-28 June 1885, Heldreich (W-HAL). Porto Rafti, May
1929, Guiol (BM). Parnes, prope Driza, Apr. 1929, Guiol (BM). Hymettus, 25 Apr.
1878, 1884, Heldreich (E ; W-HAL). Evvoia: Mt. Ocha, 23 May 1955, Rechinger
fil. 16291 (W). Peloponnisos : Messenia, May 1844, Heldreich (BM). Leontari,
May 1844, Heldreich (BM). In ruinis Olympiae, Elis, 12 Apr. 1911, Haldcsy
(W-HAL). Aegean Islands: Skyros, Apr. 1908, Tunta 663 (W-HAL). Hydra,
3-10 Apr. 1876, Heldretch (W-HAL). Melos, Mar. 1889, Heldreich & Haldcsy (BM ;
K ; W-HAL). Tenos, Apr. 1889, Heldreich & Haldcsy (K ; W-HAL). Karpathos,
Monte Lasto, 13 Mar. 1883, Pichler (BM ; K). Mytilini, Ajassos, 19 May 1934,
Rechinger fil. 5638 (W-RECH). Chios, 14-15 Apr. 1939, Platt 126 (K). Samos,
monte Kerki, 15 Apr. 1934, Rechinger fil. 4o40 (BM ; W-RECH). Kalimnos,
13 Apr. 1887, Major 676 (E).

CRETE : Viano prope Christos, May 1g00, Leonis in Dérfler goto (BM; E;
W-HAL). Hag. Vasilis, 9 May 1904, Dérfler goo (W-RECH). Lassithi, Krystal-
lenia, 23 May 1914, Gandoger 2027 (K). Kissamos, Polyrrhenia, 6 Apr. 1915,
Gandoger 8744 (K). Canee (Khania), 8 Apr. 1914, Gandoger 2792 (K).

TURKEY : Messarburnum bei Konstantinopel, 23 May 1897, Nemetz (W-RECH).
Smyrnae, 1827, Fleischer (E). Marmaris (Mugla), 24 Mar. 1956, Davis 25304 (K).
Taktali dag (Antalya), 26 May 1906, Bornmiiller gog5 (K). Sencan Deresi between
Girtimze and Siiphandere, Feke (Andana), 1 July 1952, Davis 19636 (K). Anamur
(Mersin), 14 Apr. 1956, Davis 25954 (K). Erzerum, Zorab 137 (K). Montes
Amanus, 1906, Haradjian 218 (W).

Cyprus : Kyrenia, 20 Mar. 1956, Atherton 1194 (K). West of Orga, 30 Mar.
1956, Merton 2588 (K). Above Defetra, 27 Mar. 1952, Probyn 62 (K). Larnaca,
7-20 Apr. 1912, Haradjian 28 (K). Ayios Philon, near Rizokarpeso, 19 Feb. 1941,
Davis 2257 (K). Platres, 18 June 1938, Kennedy r1gr (K). Ayia Grini (Morphan),
11 Mar. 1941, Davis 2538 (E).

U.S.S.R.: Azerbaydzhan, Plantae Bakuenses, Ins. Swatoi, 20 May 1863, Bruhns
(BM ; LIVU).

SyRIA, ISRAEL, JORDAN, LEBANON : Kishnon R., 1863-64, Lowne (K). Garnala,
3 Apr. 1911, Meyers & Dinsmore 3561 (K). Above Ain Fit, 9 May 1911, Meyers &
Dinsmore Gr561 (E ; K). Ramleh, 30 Mar. 1911, Dinsmore 1561 (E). Below
Kulaia, 31 Mar. 1877, Post 228 (K).

LisyA : Between Cyrene (Shahhat) and Apollonia (Marsa Susa), 4 Apr. 1939,
Sandwith 2350 (K).

ALGERIA : Bab-el-Oued, prés Alger, Apr. 1879, Allard, Soc. Dauph. 2388 (K).
Birmadreis, Alger, 13 Apr. and 15 May 1851, Jarmin 129 (K). Mustapha sup.,
Algiers, 10 Apr. 1873, Joad (K). Oran, 1843, anon. (K).

Introduced :

AUSTRALIA : Western Australia : Cannington, Lower Cannington River, 12 Oct.
1910, Morrison 20012 (K). Victoria: Point Lonsdale, Oct.-Nov. 1912, Tilden 672
(K). Shepparton, 17 Sept. 1925, Britten (K).

REVISION OF THE GENUS PETRORHAGIA 169

Hawall : Waimea, S. Kohala, 25 May 1938, Hosaka 2rro (K).

UNION OF SouTH AFRICA: Cape of Good Hope: Tulbagh, District Wolseley,
4 Oct. 1955, Van Breda 555 (K). Caledon to Batriver, 9 Oct. 1955, Van Breda 564
(K). Jordans Bay, Stellenbosch, 4 Oct. 1947, Parker 4268 (K).

25. Petrorhagia glumacea (Bory & Chaub.) Ball & Heywood, comb. nov.

Dianthus glumaceus Bory & Chaub. in Expéd. Sci. Morée, Sect. Sci. Phys. 3 (2): 340 (1832).
Tunica glumacea (Bory & Chaub.) Boiss., Fl. Or. 1: 517 (1867).
Kohlrauschia glumacea (Bory & Chaub.) Hayek, Prodr. Fl. Penins. Balcan. 1 : 224 (1924).
Stems up to 50 cm. high, usually simple, glabrous or scabrid. Cauline leaves up
to 40 mm. long and 2 mm. broad, linear-oblong, 3-veined ; sheath 1-2 mm. long,
usually shorter than broad. Largest bracts of head 12-18 mm. long, 8-12 mm. broad,
obtuse. Calyx (g-)10-13 mm. long; costae 3-veined ; teeth oblong, obtuse.
Petals 12-18 mm. long, the limb 3-6 mm. broad, very variable in shape, pink or
purplish. Seeds 1-5—2-2 mm. long, I-2-1-7 mm. broad, almost smooth to tuberculate.

Distribution : Balkan peninsula.

25a. Petrorhagia glumacea var. glumacea. (Fig. 22 ; Plate 15 D.)

Petal limb obdeltate, crenate to laciniate at the apex. Seeds 1-7-2:2 mm. long,
I:4-1'7 mm. broad, almost smooth or slightly tuberculate.

GREECE : Peloponnisos : Vastitza, 12-24 July 1855, Orphanides 462 (CGE ; K ;
W-HAL). Messini, May 1844, Heldreich (BM ; CGE; E). Taygetos, Likorema,
28 June 1902, Haldcsy 51 (W-HAL). Taygetos, Megali Anastasova, 1 Sept. 1899,
Heldreich (W-HAL). Prope Astros, Laconia, 6 May 1929, Guiol 556 (BM).
Laconia, Sparti, 7 June 1958, Rechinger 19984 (LIVU ; W). Carytaena (Karytaina),
21 May 1929, Guiol 1053 (BM). Plain of Argos, Apr. 1855, Mill (CGE ; K). Mycenae
(Mykene), 29 July 1888, Heider (W-HAL). Kyllene, prope Trikala, 30 June 1887,
Heldreich (W-HAL). Kalavryta, 25 June 1899, Dérfler 412 (W-HAL) ; 21 June
1926, Bornmiiller 178 (BM). Lintye (Elis), 1 June 1899, Heldreich (W-HAL). Patras,
7 June 1893, Haldcsy (W-HAL). Nea Epidavros, May 1877, Tuckett (K).
Dervenakia (Argolis), July 1930, Atchley 275 (K). Sterea Ellas: Attika, Pinatzt
(LIVU).

250. Petrorhagia glumacea var. obcordata (Marg. & Reut.) Ball & Heywood,
comb. nov.
Dianthus obcordatus Marg. & Reut. in Mém. Soc. Phys. & Hist. Nat. Genéve 8 : 281 (1839).
Kohlvauschia obcordata (Marg. & Reut.) Reichb., Ic. Fl. Germ. & Helv. 6: 43 (1844).
Tunica glumacea var. obcordata (Marg. & Reut.) Boiss., Fl. Or. 1 : 517 (1867).

Petal limb obcordate, usually entire. Seeds 1-5-1-8 mm. long, I-2-1°-5 mm.
broad, tuberculate.

GREECE : Peloponnisos : Mt. Chalkis ad sinum Patranum, 8 June 1893, Haldcsy
(K ; W-HAL). Sterea Ellas : Mt. Parnassi prope Levadeia, 17 July 1888, Haldcsy
(W-HAL). Mt. Parnassus above Delphi, 31 May 1927, Heard (BM). Gravia,
21 June 1937, Balls & Gourlay 3362 (BM; E; K). Kionae, supra Segritna,

170 REVISION OF THE GENUS PETRORHAGIA

11 July 1888, Haldcsy (W-HAL). Thessalia : Olympus, June-July 1929, Guiol
(BM). Kalampaka, 20 May 1896, Sintenis 25z (K). Orman Magula, June 1885,
Haussknecht (K). Agrapha, M. Pindi circa monast. Korona, 20-28 June 1885,
Heldreich (E). Mt. Ossa, Tsagesi, 23 July 1936, Grebenchikoff (K). Ipiros :
Metsovon, 19 July 1937, Balls & Gourlay 3649 (BM ; E; K). Mt. Mitcikeli, Sept.
1896, Baldacci (BM). Makedhonia: Kaream (Chalcidica), 1896, Dimitrijev(K). Stavros,
July 1917, Turner (K). Kefallinia : Mt. Aenos, 13 June 1895, Grimburg (W-HAL).

TuRKEY : Muratli, 20 June 1890, Degen (K).

ALBANIA : Mavrova, Valona, 27 June 1894, Baldacci 234 (BM). Biskezmi,
Ljnime, 24 Apr. 1918, Schneider (W).

JuGcosLavia : Makedonya: Roszdan, 9 July 1893, Dérfler (W-HAL). Katlan-
ovska Banja, near Skopje, seed Oct. 1953, Hort. Kew. 1954, Gore-Browne (K).
Srbya: Rissovatz, Gosnza, 3 Oct. 1889, Sintenis (K ; W-HAL). Vranja, 19109,
Lucas (BM). Bosnai Hercegovina : Uskoplje, 1 Apr. 1926, Korb (W). Crna Gora :
Borkovici, fl. Piva, July 1909, Rohlena (K). Cattaro, 13 June 1867, Huter (BM ;
CGE ; K); Puchler (BM; K; W-HAL). Hrvatska: Teodo, 4 June 1905,
Schneider (W). Krivosije, bei Ledenice, 17 June 1926, Korb (W). Scaljari, Pichler,
Fl. Austro-Hung. 2508 (K ; W-HAL ; W-RECH). Bei Combar nachst Casselnuovo,
June 1876, Studniczka (W-HAL). Circa Spalatensi, 1843, Alexander (BM).
Introduced :

ItaLty : Trentino: Prope Povo et Roncegno, July 1899, Murr in Dérfler, Herb.
Norm. 4orr (BM ; E).

This is a somewhat variable species which has been confused with P. prolifera in
the central and northern Balkan peninsula. All the Bulgarian specimens seen
and many of those from Jugoslavia and northern Greece are undoubtedly P. prolifera.

The status of the two varieties recognized is still not clear as it has not been possible
to determine the extent of the correlation between the petal and seed characters.
Most specimens are either in flower and without mature seeds or in fruit and without
petals. Large almost smooth seeds occur only in Peloponnisos and southern
Sterea Ellas while small tuberculate seeds occur in all areas except Peloponnisos.
The petal type of var. glumacea is largely restricted to Peloponnisos, but intermediate
types occur occasionally almost throughout the range of var. obcordata. A further
difficulty is that in many specimens the petals are badly pressed owing to the
bulkiness of the head so that it is difficult to see the precise shape and toothing of
the petal limb.

From this discussion it will be clear that there is the possibility that the two
varieties recognized may eventually be treated as subspecies. However, the
evidence available at the present is so inconclusive and unsatisfactory that it is
impossible to justify raising them to this rank.

Insufficiently known species
Tunica syriaca Boiss., Diagn. Pl. Or. Nov. 2 (8) : 63 (1849) ; Fl. Or. 1 : 519 (1867).—
Post, FI..Syr. Palest. & Sinai, ed. 2, 1 + 160 (1932).
“ T. glaberrima nigrescenti-viridis perennis basi suffrutescens multicaulis, caulibus

REVISION OF THE GENUS PETRORHAGIA 171

filiformibus rectis vel ascendentibus ad nodos superiores szepé geniculatis simplicibus
unifloris rarius ramulo auctis nigricantibus asperulis, foliis parvis angusté lineari-
triquetris crassiusculis obtusiusculis supra sulcatis subtus carinatis margine scabridis
strictis internodio 3-4 plo brevioribus basi breviter connatis, bracteis 8-10 calyci
subeequilongis lanceolatis acutissimis subpungentibus preter nervum rubrum
crassum angusté albo-membranaceis, calyce oblongo campanulato brevissime
puberulo tubo enervi albido dentibus tubo triplo brevioribus oblongis nigricantibus
albo-marginatis, petalis angusté linearibus obtusis ungue nudo lamina pallidé rosea
breviter exserta, capsula. . .

“Hab. in rupibus Syri@ borealis poné urbem Antiochiam sitis ubi specimina
pauca Junio 1846 legi.

‘“‘Caules tenuissimi 6-8 pollicares, folia 3-5 lin. longa, calyx cum bracteis 4 lin.
longus. Species distinctissima propé T. dianthoidem collocanda.”’

The above is a transcription of Boissier’s protologue.

Tristram (Survey of Western Palestine, Flora & Fauna : 240 (1884)) also records
this species from northern Lebanon, but he does not give any further information.
No material has been seen and no additional records have been traced. Even the
most recent description (Post, 1932) is clearly based entirely on Boissier’s data.

The description suggests that this species is nearest to Petrorhagia sect. Petrorhagia
subsect. Thessalae, all the species of which are rare and local late-flowering perennials.
Geographically it would form a link between the Balkan and North African species,
and P. macra from Kurdistan.

Excluded species

Petrorhagia glomerata (Pall. ex Bieb.) Link, Handb. Erkenn. Gewichse 2 : 235
(1831) = Gypsophila glomerata Pall. ex Bieb.

SPECIAL LITERATURE

Batt, P. W., & HEywoop, V.H. (1962). The taxonomic separation of the cytological races of
Kohlrauschia prolifera (L.) Kunth sensu lato. Watsonia 5 : 113-116.

BarkoupaH, Y. I. (1962). A revision of Gypsophila, Bolanthus, Ankyropetalum and Phryna.
Wentia 9 : 1-203.
BécHER, T. W., LarRseEN, K., & Raun, K. (1953). Experimental and cytological studies on
plant species. I. Kohlrauschia prolifera and Plantago coronopus. Hereditas 39 : 289-304.
(1955). Experimental and cytological studies on plant species. II. Tvrifolium arvense
and some other pauciennial herbs. Dan. Biol. Skr. 8 (No. 3).

BoissiER, E. (1867). Flora Ovientalis. 1. Bale & Genéve.
Danpy, J. E. (1957). Some new names in the British flora. Watsonia 4 : 41-48.
DarRLINGTON, C. D., & WyLtE, A. P. (1955). Chromosome Atlas of Flowering Plants. London.

Davis, P. H. (1957). New Turkish species of Tunica, Velezia and Potentilla. Not. R. Bot.
Gard. Edin. 22 : 163-171.

EHRENDORFER, F. (1958). Ein Variabilitaétszentrum als “ fossiler ’’ Hybrid-Komplex : Der
ost-mediterrane Galium graecum 1..-G.canum Req.-Formenkreis. Eine Monographie.
Osterr. Bot. Zeitschy. 105 : 229-279.

172 REVISION OF THE GENUS PETRORHAGIA

Hrywoop, V. H. (1958). The presentation of taxonomic information. A short guide for
contributors to Flora Europaea. Leicester.

JANCHEN, E. (1963). Catalogus Florae Austriae. I. Teil : Pteridophyten und Anthophyten
(Farne und Bliitenpflanzen). Erganzungsheft. Wien.

Léve, A., & LévE, D. (1961). Chromosome numbers of Central and Northwest European
plant species. Opera Botanica 5 : 1-581.

Marre, R. (1963). Flore del’Afrique du Nord. 10. Paris.

Wix.iams, F. N. (1890). Synopsis of the genus Tunica. Journ. of Bot. 28 : 193-199.

Bull. B.M. (N.H.) Bot. 3, 4 PLATE 13

A B

Cc D

Seeds (x15). <A, Petrorhagia illyrica subsp. haynaldiana (Janka) Ball & Heywood. B, P. lycica
(P. H. Davis) Ball & Heywood. C, P. alpina subsp. olympica (Boiss.) Ball & Heywood.
D, P. kennedyae (Jacks. & Turrill) Ball & Heywood.

Bull. B.M. (N.H.) Bot. 3, 4 PLATE 14

A B

Cc

Seeds (x15). A, Petrorhagia phthiotica (Boiss. & Heldr.) Ball & Heywood. B, P. saxifraga
(L.) Link var. saxifraga. C, P. peroninii (Boiss.) Ball & Heywood.

Bull. B.M. (N.H.) Bot. 3, 4 PLATE 15

G D

Seeds (X15). A, Petrorhagia prolifera (L.) Ball & Heywood. B, P. nanteuilit (Burnat) Ball &
Heywood. C, P. velutina (Guss.) Ball & Heywood. D, P. glumacea (Bory & Chaub.) Ball &
Heywood var. glumacea.

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MARINE ALGAE OF GOUGH
ISLAND

YVONNE M. CHAMBERLAIN

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 3 No. 5
LONDON : 1965

fen
a
hee oa,

MARINE ALGAE OF GOUGH ISLAND

BY

YVONNE M. CHAMBERLAIN

Pp. 173-232 ; 80 Text-figures ; Plates 16-19

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 3 No. 5
LONDON : 1965

THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, 1s
issued in five series corresponding to the Departments
of the Museum, and an Historical series.

Parts will appear at irregular intervals as they become
veady. Volumes will contain about three or four
hundred pages, and will not necessarily be completed
within one calendar year.

This paper ts Vol. 3 No. 5 of the Botany serves.

© Trustees of the British Museum (Natural History), 1965

TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)

Issued May, 1965 Price Thirty-two Shillings

MARINE ALGAE OF GOUGH ISLAND

By YVONNE M. CHAMBERLAIN
(Mrs. BUTLER.)

THE material reported on in this paper was collected by Dr. N. M. Wace during the
stay of the Gough Island Scientific Survey on Gough Island from 13 November
1955 to 13 May 1956, and is now in the British Museum Herbarium. The collections
were made on only a few days during this summer period and the species included
here undoubtedly represent only a few of those present on the island. However,
as Dr. Wace is an ecologist, he collected particularly those species which dominated
the littoral and sub-littoral zones ; I think therefore that all the zone-forming
species are included here together with the most abundant associated littoral
species. Only two or three attempts at dredging were made and these were
unfortunately fruitless ; none, therefore, of the smaller sub-littoral species, notably
the Delesseriaceae, which undoubtedly grow round Gough Island, is included in the
collections.

ACKNOWLEDGEMENTS

I am indebted to Dr. Wace for making the collections, which were well preserved
and clearly labelled, also to him and Dr. M. W. Holdgate (the invertebrate zoologist
of the expedition) for much helpful discussion on the littoral ecology of the island,
and to Mr. J. J. van der Merwe who collected some more Blidingia and Enteromorpha
when he stayed on Gough Island after the return of the Survey and thus helped to
clarify some taxonomic and ecological problems.

I worked on the material while I was a member of the staff of the British Museum
(Natural History) and I am grateful to Mrs. F. L. Balfour-Browne, Mr. J. E. Dandy
and Mr. R. Ross for general, editorial and nomenclatural help ; to Dr. W. T. Stearn
who wrote the latin diagnoses of new species ; and to Mr. D. W. Cooper who made
sections of the Corallinaceae.

For general taxonomic guidance I am indebted to Dr. P. S. Dixon and to Mme.
Marie Lemoine, with whom I was able to discuss the Corallinaceae when I visited
the Muséum National d’Histoire Naturelle, Paris. Several of the species were sent
to taxonomists expert in particular groups and their help is acknowledged in the
appropriate places.

Finally, I would like to thank the Directors of the following Institutions for
permission to examine herbarium material: Royal Botanic Gardens, Kew ;
Bolus Herbarium, Cape Town ; Herbarium of the University of California ;
Rijksherbarium, Leyden ; Muséum National d’Histoire Naturelle, Paris ; and
Dr. Egil Baardseth, who presented a duplicate collection of his Tristan da Cunha
algae to the British Museum.

MARINE ALGAE OF GOUGHIISLAND

GEOGRAPHY OF GOUGH ISLAND

Gough Island (Figs. 1, 2) is an outlier of the Tristan da Cunha group of islands
situated near the southern end of the mid-Atlantic ridge. It lies 370 km. S.S.E.
of Tristan da Cunha, more than 3,000 km. from South America to the west and

Greenwich

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Fic. 1. The Southern Hemisphere.

2,400 km. from South Africa to the north-east. The island lies in the West Wind
Drift and within the zone of sub-antarctic, cold-temperate water (Knox, 1960, p. 578)

but may occasionally be influenced by the cold-temperate mixed water derived from
the Brazil current. Gough Island lies between the Antarctic and Sub-tropical
Convergences (Knox, 1960, p. 582) but is closer to the latter, which is here at, or

somewhat to the south of, Tristan da Cunha.

MARINE ALGAE OF GOUGH ISLAND 177

Church
i] Rock
0

Point Dell Rocks
Window Rock

Slaughtered
Seal Bay

“o
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Midshipman

» Stand off
Rock

[000 yds_

Fic. 2. Sketch map of Gough Island to show localities ee marine algae were collected.
LITTORAL ECOLOGY OF GOUGH ISLAND

It is intended to publish a paper on the littoral ecology of Gough Island and only
the main features will be summarized here.

The entire island is very exposed in comparison with continental shores so that
the terms sheltered, moderately exposed, etc., are relative only to conditions on
the island. On the moderately exposed shore at Dell Rocks Beach the zonation is :

Upper supra-littoral Lichina sp.
Lower supra-littoral Porphyra tristanensis, Verrucaria sp.

Upper littoral Blidingia minima in crannies and sheltered places.
Inidaea laminarioides where more exposed. Enteromorpha
bulbosa.

Mid littoral Inidaea laminarioides gradually giving way to Rhodo-
glossum revolutum.

Lower littoral Inridaea laminarioides and Rhodoglossum revolutum
succeeded by Durvillaea antarctica and Melobesteae.

Sub-littoral The Durvillaea-Melobesieae zone extends down into the

sub-littoral and below this Macrocystis pyrifera is
dominant.

178 MARINE ALGAE OF GOUGH ISLAND

Plate 16 A shows the shore at Isolda Rock which is somewhat more exposed than
Dell Rocks Beach. Light-coloured Melobesteae can be seen on the left, a dark band
of Iridaea laminariotdes in the centre and plants of Durvillaea antarctica on the right.

On the more sheltered Midshipman Rock, the mid littoral is dominated by a band
of Ralfsia sp., below which is a band of Melobesieae and Corallina officinalis. West
Point Reefs is the most exposed locality ; here Ividaea laminarioides and Rhodo-
glossum revolutum zones are absent and the mid littoral is dominated by a band of
white Melobesteae succeeded by a band of Polysitphonia howei (probably) and other
Rhodomelaceae. Macrocystis pyrifera is also absent on very exposed shores.

Among the most common plants occurring in littoral rock pools throughout the
island are: Corallina officinalis, Dermatolithon nodulosum, Inidaea undulosa,
Scytostphon lomentaria and Codium fragile.

The algal zonation on Gough Island is typical of the sub-antarctic pattern
(Skottsberg, 1941 ; Knox, 1960, p. 601). It differs markedly from that of Tristan
da Cunha, which has more in common with a South African shore. This corresponds
with the fact that Gough Island is well south of the Sub-tropical Convergence while
Tristan da Cunha is at its mean position. Very little is known about the east
coast shore of South America at about the same latitude as Gough Island, but it is
probable that further investigations will show that it has a zonation pattern similar
to that of Gough Island. Further south, Tierra del Fuego and the Falkland Islands
have a zonation pattern similar to that of Gough Island but with a greater number
of typical antarctic species.

LOCALITIES AND HABITATS, WITH SPECIES PRESENT

(Entries in brackets relate to algae that have not been identified at species level
and are not included in the Systematic List.)

Dell Rocks Point. 10 Dec. 1955. Washed ashore at storm-tide mark. G.J.S.S.
1501. {Melobesieae.|

Between the Admiral and the Commodore. 22 Dec. 1955. Between 18 and
55 m., and in gullies. G.J.S.S. 1502. Macrocystis pyrifera.

Slaughtered Seal Bay. 22 Dec. 1955. Porphyra zone. G.I.S.S. 1503. Porphyra
tristanensis.

Slaughtered Seal Bay. 22 Dec. 1955. In pools from water level down to Io cm.
G.I.S.S. 1504. Ulva lactuca, Corallina officinalis, Dermatolithon nodulosum.

Slaughtered Seal Bay. 22 Dec. 1955. Below15cm.inrock pools. G.I.S.S. 1505-
1507. Codium fragile with epiphytic Feldmannia globifera, Giffordia granulosa and
Sphacelaria furcigera, Gelidium regulare, Corallina officinalis, Dermatolithon
nodulosum, Lithothamnium neglectum, Iridaea undulosa, Centroceras clavulatum.

Beach south of Sophora Glen. 28 Dec. 1955. At tide mark on boulder beach.
G.I.S.S. 1508. |Melobesieae, Gelidium sp.]

Near Penguin Island. 2 Jan. 1956. At55m. G.I.S.S. 1509. Macrocystis pyrifera,
[Melobesieae].

Outer Dell Rocks. 18 Feb. 1956. From mid-tide level to 2 m. below low-tide level.
G.I.S.S. 1510. Durvillaea antarctica.

MARINE ALGAE OF GOUGH ISLAND 179

Dell Rocks Beach. 19 Feb. 1956. Near low-tide mark on boulder beach. G.J.S.S.
511. Scytosiphon lomentaria.

Window Rock. 18 Feb. 1956. Below low-water neaps on rocks amongst corallines.
G.I.S.S. 1512. Cladophora radtosa.

Midshipman Rock. 18 Feb. 1956. Porphyra zone, upper shore. G.I.S.S. 1573,
1514. Oscillatoria nigroviridis, Phormidium submembranaceum, Ulva lactuca,
Porphyra tristanensis, Dermatolithon nodulosum, Iridaea laminarioides, Rhodo-
glossum revolutum.

Midshipman Rock. 18 Feb. 1956. Mid-littoral zone, on shiny olive-green deposit.
G.I.S.S. 1515, 1517, 1518. [Ralfsia|, Corallina officinalis, Centroceras clavulatum,
Microcladia alternata, |Delesseriaceae|, Herposiphonia paniculata, Polysiphonia
howet.

Midshipman Rock. 18 Feb. 1956. Lower littoral. GJ.S.S. 1516. Colpomenia
sinuosa.

Midshipman Rock. 18 Feb. 1956. Standing water in upper littoral. G.J.S.S.
I519, 1520. Scytosiphon lomentaria, Centroceras clavulatum, Herposiphonia
paniculata, | Rhodomelaceae}.

Midshipman Rock. 18 Feb. 1956. Upper-littoral pools. G.J.S.S. 1527, 1522.
Entophysalis conferta, E. deusta, Calothrix crustacea, Lyngbya confervoides, Ulva
lactuca.

Standoff Rock. 18 Feb. 1956. Mid-littoral coralline zone. G.J.S.S. 1524, 1525.
Corallina officinalis, Rhodoglossum revolutum.

Standoff Rock. 18 Feb. 1956. Above Corallina on less-exposed part of rock.
G.I.S.S. 1526. Rhizoclonium ambiguum, R. riparium forma validum, Corallina
officinalis, Iridaea laminarioides, Bostrychia mixta, Lophostphonia scopulorum.

Standoff Rock. 18 Feb. 1956. Upper littoral, Porphyra zone. G.I.S.S. 1527.
Entophysalis deusta, Calothrix crustacea, Plectonema calothrichoides, Bangia
fuscopurpurea, Porphyra tristanensis, |Hildenbrandia}.

Church Rock. 5 Mar. 1956. Vertical rock face. G.J.S.S. 1528-1535. Cladophora
vadiosa, Codium adhaerens, Iridaea undulosa, Rhodoglossum revolutum, Centroceras
clavulatum, |Nitophyllum sp.], Herposiphonia paniculata, Lophurella sp.

West Point Reefs. 6 Mar. 1956. In and around bird-fouled spray-zone rock pools.
G.I.S.S. 1536-1538, 1544. Entophysalis deusta, Calothrix crustacea, Plectonema
calothrichoides, Enteromorpha bulbosa, Scytosiphon lomentaria, |Hildenbrandia? |.

West Point Reefs. 6 Mar. 1956. In rock pools below spray zone. G.I.S.S. 1540,
1541. Ulva lactuca, Codium fragile, Corallina officinalis, Dermatolithon nodulosum,
Rhodoglossum revolutum, Polysiphonia boergesenit.

West Point Reefs. 6 Mar. 1956. Margins of pools below spray zone. G/.S.S.
1542, 1543. Ulva lactuca, Dermatolithon nodulosum.

Isolda Rock. 6 Mar. 1956. Mid-littoral rock pools. G.I.S.S. 1545, 1546, 1550,
1551. Cladophora rupestris, Corallina officinalis, Dermatolithon nodulosum,
Inridaea undulosa.

Isolda Rock. 6 Mar. 1956. Lower-littoral rock pools. G.J.S.S. 1547, 1548. Cor-
allina goughensis, Lithothamnium neglectum.

180 MARINE ALGAE OF GOUGH ISLAND

Isolda Rock. 6 Mar. 1956. Margins of mid-littoral pools. G.J.S.S. 1549. Poly-
siphonia howe.

Isolda Rock. 6 Mar. 1956. Upper littoral on ridges between pools. G.J.S.S.
1552. Iridaea laminarioides, Rhodoglossum revolutum, Herposiphonia paniculata,
Polysiphonia hower.

Isolda Rock. 6 Mar. 1956. Upper-littoral pools. GJ.S.S. 1553. Corallina
officinalis, Dermatolithon nodulosum, Lithothamnium neglectum, Rhodoglossum
revolutum, Centroceras clavulatum.

Window Pool. 12 Apr. 1956. In caves and wet crevices of spray-zone rocks.
G.I.S.S. 1554. [Chlorophyta].

Window Pool. 12 Apr. 1956. In pools and sheltered channels of boulder beach.
G.I.S.S.1555,1556. Blidingia minima, Inidaea undulosa, Rhodoglossum revolutum.

Window Pool. 12 Apr. 1956. On walls of small shallow cave beside pool. G.I.S.S.
1557. Entophysalis conferta, Rhizoclonium ambiguum.

Dell Rocks Beach. 12 Apr. 1956. In sheltered places between boulders and in
shallow pools. G.I.S.S. 1558. Guiffordia mitchelliae.

Dell Rocks Beach. 12 Apr. 1956. In pools and protected channels just above
mid-tide level. G.I.S.S. 1559. Scytostphon lomentaria.

Between Station and Dell Rocks. 1 Dec. 1960. High-water level. Van der
Merwe 1,2, 4. Blidingia minima, Enteromorpha bulbosa.

Capsize Sands. 1 Dec. 1960. Van der Merwe 3. Blidingia minima, Enteromorpha
bulbosa.

PHYTOGEOGRAPHY

The elements of which the Gough Island marine algal flora is composed are :

Cosmopolitan 19 species
General southern hemisphere 4 species
Southern cold-temperate 6 species
North and south Atlantic 3 species
Tristan da Cunha group endemics 6 species
Endemic 2 species

This list is necessarily incomplete and inaccurate for two reasons. The first is
that, compared with, for example, the angiosperms, relatively little is known
about the distribution of marine algae and, until fairly recently, authors have made
little attempt to relate the flora of the area on which they were working to that of
other areas. Secondly, there is at present a very wide variation of opinion among
algal taxonomists as to what constitutes a species. In the past, a large number of
new species have been described without regard to seasonal, ecological or geo-
graphical variation, and also without reference to previous literature on or collections
from the same area. On the whole, present-day phycologists regard species in the
widest sense, as entities which are liable to great variation according to the external
conditions mentioned above. I agree with this concept, and this may account for
the large number of cosmopolitan and widely distributed species in the list.

MARINE ALGAE OF GOUGH ISLAND 181

It is a well-known fact, however, that isolated islands usually have relatively
high numbers of two groups of species in their flora, cosmopolitan species and endemic
species, so that the list may be only an exaggeration of the true picture. Of the
second group, the endemic species, two species so far known only from Gough
Island are described, and as these are in the Corallinaceae, which is a very difficult
and often ignored family, it is probable that they will later be found elsewhere.
On the other hand, endemic species are more likely to be found among the smaller,
less conspicuous species. As previously explained, the collector concentrated on
the dominant species and many possible endemics have no doubt so far escaped
notice. There were some more possibly new species in the collection but the
material was too scanty, in my opinion, to merit their description as such.

There are some rather surprising absences from Gough Island, most notably
Hildenbrandia lecannelliert and Adenocystis utricularis. Sub-littoral members of the
Delessertaseae and of Desmarestia are also missing from the collection, probably
because of the difficulties of dredging. Two littoral members of the Delesseriaceae
were collected but I was unable to identify them further than that they are
probably species of Nitophyllum. The Hildenbrandia and Adenocystis may have
been missed, but Dr. Holdgate, who has seen many southern cold-temperate
shores, is fairly sure that they were not at any of the localities visited by the Survey.

In conclusion, on present evidence it is not possible to make many generalizations
about the Gough Island marine algal flora. It does however represent a typical sub-
antarctic zonation pattern and the high number of endemic species (to the Tristan
da Cunha group) is typical of isolated localities.

SYSTEMATIC LIST
CYANOPHYTA!

CHAMAESIPHONACEAE
ENTOPHYSALIS CONFERTA (Kiitz.) Drouet & Daily in Lloydia 11 : 79 (1948) ; in
Butler Univ. Bot. Stud. 12 : 111, figs. 196-215 (1956).
Palmella conferta Kiitz., Phyc. Germ.: 149 (1845).

Midshipman Rock, in upper-littoral pool, 18 Feb. 1956, G.I.S.S. 1522. Window
Pool, on walls of small shallow cave beside pool, 12 Apr. 1956, G.I.S.S. 1557.

Cosmopolitan.

This species occurred as an epiphyte on Calothrix crustacea and Rhizoclonium
ambiguum. When Dr. Drouet discussed the identification of the Gough Island
Cyanophyta with me, he commented that the only difference between this and the
next species is that this is an epiphyte and E. deusta is a lithophyte.
ENTOPHYSALIS DEUSTA (Menegh.) Drouet & Daily in Lloydia 11 : 79 (1948) ; in

Butler Univ. Bot. Stud. 12 : 103, figs. 185-194, 247-250 (1956).
Coccochloris deusta Menegh. in Atti 2 Riun. Sci. Ital. Torino 1840 : 173 (1841).

1Dr. Francis Drouet of the Academy of Natural Sciences of Philadelphia kindly identified the
Cyanophyta.

182 MARINE ALGAE OF GOUGH ISLAND

Midshipman Rock, in upper-littoral pool, 18 Feb. 1956, G.I.S.S. 1522. Standoff
Rock, from upper littoral, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1527. West Point
Reefs, encrusting layer on rock in and around spray-zone pools, 6 Mar. 1956,
G.I.S.S. 1538, 1544.

Cosmopolitan.

The growth form of the specimens of this plant from Gough Island is the cushion
type. It grows on rocks in the upper littoral zone where it forms a film, usually in
association with other Cyanophyta. Calothrix crustacea is its most frequent associate.

RIVULARIACEAE

CALOTHRIX CRUSTACEA Born. & Flah. in Ann. Sci. Nat., Sér. 7, Bot. 3 : 359 (1886).
Midshipman Rock, in upper-littoral pool, 18 Feb. 1956, G.J.S.S. 1522. Standoff
Rock, from upper littoral, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1527. West Point
Reefs, encrusting layer on rock in and around spray-zone pools, 6 Mar. 1956,
GIS3S 7538; 1544:
Cosmopolitan.

C. crustacea is the most common blue-green alga collected on Gough Island. It
grows in pools and on rocks in the upper littoral and spray zones of several localities,
where it forms a film mixed with Entophysalis deusta and Lyngbya confervoides. It
often bears epiphytes such as Plectonema calothrichoides and Entophysalis conferta.
From the available information, there is no indication that the Cyanophyta are ever
the dominants of a littoral zone on Gough Island, although they are certainly a
noticeable feature of the upper littoral zone on the Midshipman Rock. Baardseth
(I94I, p. 133) records Cyanophyta as forming a dense covering of rocks in the
Porphyra zone on the exposed rocks of Tristan da Cunha and they frequently occur
as the dominant or sub-dominant algal community at this shore level in other parts
of the world. Skottsberg (1941) did not mention the Cyanophyta in his analysis of
antarctic and sub-antarctic algal communities.

OSCILLATORIACEAE

LYNGBYA CONFERVOIDES Gom. in Ann. Sci. Nat., Sér. 7, Bot. 16 : 136, t. 3 figs. 5, 6
(1892).
Midshipman Rock, in upper-littoral pool, 18 Feb. 1956, G.I.S.S. 1522.
Cosmopolitan.

Collected from the Midshipman Rock where it grew with Calothrix crustacea and
Entophysalis deusta as a film on rock submerged in an upper-littoral rock pool.

OSCILLATORIA NIGROVIRIDIS Gom. in Ann. Sci. Nat., Sér. 7, Bot. 16: 217, t. 6
fig. 20 (1892).
Midshipman Rock, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1514.
Cosmopolitan.

This species was found growing with Phormidium submembranaceum on the
Midshipman Rock.

MARINE ALGAE OF GOUGH ISLAND 183

PHORMIDIUM SUBMEMBRANACEUM Gom. in Ann. Sci. Nat., Sér. 7, Bot. 16: 180,
t. 5 fig. 13 (1892).
Midshipman Rock, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1514.
Cosmopolitan.

Only a small amount of this alga was collected ; it came from the upper littoral
zone on the Midshipman Rock where it was growing mixed with Oscillatoria
nigroviridis forming a film on the rocks.

PLECTONEMA CALOTHRICHOIDES Gom. in Bull. Soc. Bot. France 46 : 30, t. 1 figs. 6-10
(1899).

Standoff Rock, from upper littoral, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1527.
West Point Reefs, encrusting layer on rock in and around spray-zone pools, 6 Mar.
1956, G.I.S.S. 1544.

Cosmopolitan.

Growing as an epiphyte on Calothrix crustacea.

CHLOROPHYTA
ULVACEAE

BLIDINGIA MINIMA (Nageli ex Kiitz.) Kylin in K. Fysiogr. Sallsk. Lund Forhandl.
17: 181 (1947).—Womersley in Austral. Journ. Mar. Freshw. Res. 7: 350

(1956). (Figs. 3-5.)

Ulva intestinalis var. nana Sommerfelt, Suppl. Fl. Lapp.: 186 (1826).
Enteromorpha minima Nageli ex Kiitz., Sp. Alg.: 482 (1849).—Kiitz., Tab. Phyc. 6: 16,
t. 43 fig. 3 (1856).—Bliding in Bot. Notis. 1938: 84, figs. 1-6 (1938).—Sj6stedt in
Svensk Bot. Tidskr. 33 : 18, fig. 1 (1939).
Enteromorpha micrococca Kiitz., Tab. Phyc. 6 : 11, t. 30 fig. 2 (1856).—T. A. Stephenson in
Ann. Natal Mus. 11 : 296 (1948).
Enteromorpha gunniana J. G. Agardh in Act. Univ. Lund. 19 (2) : 122 (1883).—Hylm6 in
K. Svensk. Vetenskapsakad. Handl, Ser.. 3, 17 (1) : 6 (1938).—Sjéstedt, tom. cit. : 54,
fig. 11 (1939).—Skottsb. in K. Svensk. Vetenskapsakad. Handl., Ser. 3, 19 (4) : 75 (1941).
—Levring in Act. Univ. Lund., Ny Foljd, Avd. 2, 56 (10) : 8 (1960).
Enteromorpha nana (Sommerfelt) Sjéstedt, tom. cit.: 28, figs. 3-6 (1939).—V. J. Chapman
in Journ. Linn. Soc. Lond., Bot. 55 : 412, figs. 57-63 (1956).
Window Pool, abundant in pools and sheltered channels of boulder beach, 12 Apr.
1956, G.I.S.S. 1555. Between Station and Dell Rocks, high-water level, 1 Dec. 1960,
Van der Merwe 2. Capsize Sands, 1 Dec. 1960, Van der Merwe 3.

Cosmopolitan.

This plant grew abundantly in the upper and mid littoral zones at Dell Rocks
Beach and occurred both in sheltered rock crevices and in pools. It was also
collected from Capsize Sands. It formed a low, bright-green turf which at times
dominated the shore community. This species is found throughout the world in the
upper littoral zone of the sea-shore and quite often dominates parts of this zone
(cf. T. A. & A. Stephenson, 1954, p. 38).

The Gough Island material consists of minute plants rarely more than 1 cm. high.
Numerous fronds grow from a flattened basal part (Fig. 3) ; the fronds are tubular,

184 MARINE ALGAE OF GOUGH ISLAND

~

Imm

Fics. 3-5. Blidingia minima (Nageli ex KUtz.) Kylin: 3, habit of plant ; 4, surface
view of frond, = pyrenoid ; 5, transverse section of frond, os = outer surface, p =
pyrenoid.

the tubes being closed when young but open at the top when older. Some of the
older plants are twisted while others are swollen at the top. The cells seen in surface
view are irregular in shape and arrangement (Fig. 4) ; they have thick walls and a
large single pyrenoid. The cells measure up to 7u in diameter but are mostly about
3-5u. In transverse section (Fig. 5) the cells are elongated and measure about 3-4
x 8-gu, and the whole section is about 15-18 deep. The inner wall is thick and
may measure up to 8u deep.

Numerous forms and varieties of Blidingia minima have been described, under the
various names given in synonymy above, according to the size and shape of the fronds,
the thickness of the cell walls and the nature of the basal part. The Gough Island
material is most like the descriptions of Enteromorpha gunmniana (e.g. that of
Sjdstedt), but Womersley, who has seen Agardh’s type material, does not think it
sufficiently distinct to merit specific separation from the rest of the complex.

Dr. Carl Bliding kindly confirmed the identification of this species.

ENTEROMORPHA BULBOSA (Suhr) Montagne in Gaudich., Voy. Monde La Bonité,
Bot. 1 : 3 (1846).—De Toni, Syll. Alg. 1 : 127 (1889). (Figs. 6-8.)

Solenia bulbosa Suhr in Flora 22 : 72, t. 4 fig. 46 (1839).

Enteromorpha africana Kiitz., Phyc. Gen. : 300 (1843) ; Tab. Phyc. 6: 14, t. 40 fig. 2
(1856).

Enteromorpha hookeriana Kiitz., Sp. Alg.: 480 (1849) ; Tab. Phyc. 6: 13, t. 37 fig. 2
(1856).

West Point Reefs, in bird-fouled spray-zone rock pools, 6 Mar. 1956, G.I.S.S. 1537.
Between Station and Dell Rocks, high-water level, 1 Dec. 1960, Van der Merwe 1 ;
same locality, seaweed turning white in sun, 1 Dec. 1960, Van der Merwe 4. Capsize
Sands, 1 Dec. 1960, Van der Merwe 3.

South America, Falkland Islands, South Africa, Crozet Islands, Kerguelen Island,
Australia, New Zealand, Chatham Islands, Antarctica, ? Japan.

MARINE ALGAE OF GOUGH ISLAND 185

Collections of E. bulbosa were made at the very exposed West Point Reefs where
it was growing in penguin-fouled rock pools in the spray zone, at Dell Rocks Beach
where it grew on littoral rocks, and at Capsize Sands. The green band recorded as
being present on other parts of the island was probably dominated by this species and
Blidingia minima.

Fics. 6-8. Enteromorpha bulbosa (Suhr) Montagne : 6, part of mature plant ; 7, surface
view of frond, p = pyrenoid ; 8, transverse section of frond, os = outer surface.

The largest plants of this species collected from Gough Island (Fig. 6) are
6 cm. high and up to 5 mm. wide, but most of the plants are smaller than this. Both
branched and unbranched plants were collected ; when branches are present they
are long and very similar to the main axis, as is characteristic of this species, and not
noticeably smaller than the main axis as in many other species of Enteromorpha.
The thallus is tubular and compressed, the branches and axis are usually open at the
top and often torn. The branches and main axis taper downwards to a terete base
and some specimens still have the warty basal part by which they had been attached
to the rocks. Many of the tubes are perforated (Fig. 6) by an amphipod ; this
animal apparently lives in the tubes of FE. bulbosa and many specimens of it were
found in situ.

When seen in surface view (Fig. 7) the cells are irregularly arranged, are up to
I5u in diameter, and have rounded corners. There is one fairly large pyrenoid per
cell. In transverse section (Fig. 8) the cells are rectangular and measure about
25u. tall x 124 wide. The inner and outer membranes of the tube are fairly thick

186 MARINE ALGAE OF GOUGH ISLAND

and the section measures about 30u deep altogether. The plants are grass-green in
colour and tend to be darker at the base of the axis and branches than in the upper
parts, a feature mentioned by von Suhr.

The Gough Island plants agree well with von Suhr’s description, with Montagne’s
material in the herbarium of the Muséum National d’Histoire Naturelle, Paris, and
with Kiitzing’s figures of E. africana and E. hookeriana, both considered by De Toni
to be E. bulbosa.

E. bulbosa is a common plant of the sub-antarctic littoral ; according to
Skottsberg (1941, p. 56) it forms a widespread community on shores with a loose,
mobile substrate such as sand, and frequently occurs also in other associations such
as littoral pools. £. bulbosa has been recorded from the antarctic (cf. Hariot, 1907,
p. 3, and Gain, 1912, p. 26) and from warmer parts such as South Africa (Levring,
1938, p. 5), New Zealand (Chapman, 1956, p. 411) and Australia (Womersley, 1956,
Pp. 351), but it is on sub-antarctic shores that it grows most prolifically.

The identification of this species was confirmed by Dr. Bliding.

DivatrActuca L., Sp: Pl2- 1103 (1753)e0 (Pigs. 6-12.)

Slaughtered Seal Bay, in pools from water level down to I0 cm., 22 Dec. 1955,
G.I.S.S. 1504. Midshipman Rock, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1514 ;
same locality, dominant in upper-littoral pools, 18 Feb. 1956, G.J.S.S. 1527. West
Point Reefs, pools below spray zone, 6 Mar. 1956, G.I.S.S. 1541 ; same locality,
margins of pools below spray zone, 6 Mar. 1956, G.I.S.S. 1542, 1543.

Cosmopolitan.

This plant was very common on rocks and in pools in the upper littoral zone of the
Gough Island shores.

{RG EP ST ba

( Bi6 10u
10 il

iz

Fics. 9-12. UlvalactucaL. : 9, blade margin ; 10, surface view of frond ; 11, transverse
section of frond ; 12, transverse section of stipe, semi-diagrammatic.

The material collected from Gough Island varies considerably in size and shape.
All the specimens have a somewhat fleshy stipe from which the blades expand ;
the blades are sometimes entire but more often divided—sometimes right down to

MARINE ALGAE OF GOUGH ISLAND 187

the holdfast. The blade margins (Fig. 9) vary from smooth to crenulate or
fimbriate, all types of margin sometimes occurring on one plant. Although there is
considerable variation in external appearance, the microscopic structure of the plants
is remarkably constant. In the upper parts of the plants the cells are irregularly
arranged, somewhat rounded and up to 20yu in diameter in surface view (Fig. 10).
In section the blade is 60—70u thick (Fig. 11), the two rows of cells are separated by
a hyphal layer and the cells are rather rounded and not more than one and a half
times as high as they are wide. The stipe in section (Fig. 12) is much thicker than
the blade and has a wide central region of hyphae.

This material undoubtedly corresponds to what has been generally accepted as
Ulva lactuca L.—an extremely common plant which grows throughout the world.
Papenfuss (1960, pp. 303-305) has investigated Linnaeus’s material of U. lactuca
and his concept of the genus Ulva. As a result of this investigation, Papenfuss
concluded that the genus which Linnaeus really had in mind was that now known as
Enteromorpha Link and proposed that the generic name Ulva should be conserved
in the sense of Thuret and all subsequent authors ; this proposal has now been
accepted, and I am here using the name in that sense.

CLADOPHORACEAE

RHIZOCLONIUM AMBIGUUM (Hook. & Harv.) Kiitz., Sp. Alg. : 387 (1849). (Figs.

13-15.)

Conferva ambigua Hook. & Harv. in Hook., Lond. Journ. Bot. 4 : 295 (1845).

Rhizoclonium hookeri Kiitz., op. cit. : 383 (1849) ; Tab. Phyc. 3 : 21, t. 67 fig. 3 (1853).—
Stockmayer in Verh. Zool.-bot. Ges. Wien 40 : 584 (1890).—V. J. Chapman in Journ.
Linn. Soc. Lond., Bot. 55 : 468, fig. 126 a (1956).—W. R. Taylor, Mar. Alg. E. Trop. &
Subtrop. Coasts Amer. : 77, t. 2 fig. 5 (1960).

Rhizoclonium africanum Kiitz., Tab. Phyc. 3 : 21, t. 67 fig. 2 (1853).

Lychaete novae-zelandiae J. G. Agardh in Act. Univ. Lund. 14 (4) : 2 (1878).

Rama novae-zelandiae (J. G. Agardh) V. J. Chapman in Trans. R. Soc. New Zeal. 80:
56, fig. 11 (1952).

Standoff Rock, on less-exposed part of the rock above the Corallina zone, 18 Feb.
1956, G.I.S.S. 1526. Window Pool, on walls of small shallow cave beside pool,
¥2 Apr. 1950,G.1 58:5. 1557:

Widely distributed in the Southern Hemisphere and the tropics
as far north as Florida.

; recorded from

A small quantity of this species was found in the Bostrychietum on Standoff Rock ;
it occurred in greater quantity in a shallow cave beside Window Pool. The latter
habitat compares with the grottos mentioned by Taylor.

The plants consist of sparingly branched filaments (Figs. 13-15), the cells of
which measure 60—goy wide and up to 150u long. The cell walls are thick and
lamellated. Rhizoidal outgrowths are quite frequent and they usually terminate
in a digitate hapteron which often attaches itself to other filaments of the plant. The
cell contents do not show much structure, but prominent, dark, parietal bodies are
present, which are waste products. A few sporangia are present but the nature of
the spores cannot be determined in this preserved material.

188 MARINE ALGAE OF GOUGH ISLAND

1Ou

i TT a a a)
lO
15

Waa ala:

Fics. 13-15. Rhizoclonium ambiguum (Hook. & Harv.) Kiitz. : 13, branched filament ;
14, filaments with rhizoid, y = rhizoid, h = hapteron, w = waste products ; 15, detail
of branching.

The Gough Island plants are very similar to the type material in the Kew
Herbarium and the isotype material in the British Museum Herbarium. Stockmayer
was of the opinion that Kiitzing’s Rhizoclonium hookeri and R. africanum were to be
referred to the same species ; this view is now generally accepted. The type of R.
hookeri is a specimen collected by J. D. Hooker on Kerguelen Island at the same time

MARINE ALGAE OF GOUGH ISLAND 189

as he collected his Conferva ambigua but which he labelled C. linum. There is an
iostype of R. hooker: in the British Museum Herbarium which in all ways exactly
resembles R. ambiguum. This species is often called R. hookert by modern authors,
but if, as most authors agree, that species and Rk. ambiguum are to be united, the
latter name is correct because the publication of its basionym antedates that of
R. hookert.

The characteristic features of R. ambiguum are its sparse branching—most species
of Rhizoclonium are unbranched—and the filament width, which ranges from
40u to 100u. It is usually found in marine habitats but is also recorded from
brackish and fresh water (Taylor, loc. cit.). The type specimen was marine, although
Kiitzing incorrectly stated that it came from fresh water. In 1952 Chapman
erected a new genus, Rama, which differed from Rhizoclonium only in being sparingly
branched ; to this genus he transferred Lychaete novae-zelandiae. Chapman said
that Rama was similar to Spongomorpha. However, as the branching in Rama is
dichotomous (Figs. 13, 15) as opposed to the characteristic lateral branching of
Spongomorpha, they are not really very similar. In 1956 Chapman repeated his
description of Rama novae-zelandiae and treated. Rhizoclonium hookeri, which he
described as being unbranched and 47-70 wide, as a separate species, retaining it
in the genus Riizoclonium. In Bornet & Thuret’s herbarium (in the Muséum National
d’Histoire Naturalle, Paris) there is an isotype of Lychaete novae-zelandiae collected
by Berggren at Warrington, New Zealand. This plant is sparingly branched and has
rhizoids, the cell walls are thick and lamellated, and the filaments measure 60—80y.
wide. It closely resembles the type material of R. ambiguum. In my view, Rama
novae-zelandiae as well as Rhizoclonium hookert should be united with R. ambiguum.
This species is characteristically branched, but this feature alone does not seem
sufficient basis for generic separation and I have retained it in the genus Rhizoclonium.

RHIZOCLONIUM RIPARIUM (Roth) Harv., Phyc. Brit.: t. 238 (1849).
Forma VALIDUM (Hansg.) Foslie in Tromso Mus. Aarshefter 13 : 138 (1890).—
Koster in Pubbl. Staz. Zool. Napoli 27 : 351 (1955). (Figs. 16-18.)
Conferva tortuosa Dillw., Synops. Brit. Conferv.: t. 46 (1805)?.
Rhizoclonium tortuosum (Dillw.) Kiitz., Phyc. Germ.: 205 (1845).—V. J. Chapman in
Rhodora 41-: 20 (1939).
Rhizoclonium riparium [subsp.?] b) validum Hansg. in Arch. Naturw. Landesdurchforsch.
Bohm. 5 (6) : 79 (1886).
Rhizoclonium riparium var. validum (Hansg.) De Toni, Syll. Alg. 1 : 279 (1889).
Lola tortuosa (Dillw.) V. J. Chapman in Journ. Linn. Soc. Lond., Bot. 55 : 463 (1956).

Standoff Rock, on less-exposed part of the rock above the Corallina zone, 18 Feb.
1956, G.I.S.S. 1526.
Probably cosmopolitan.

This alga was collected from the upper littoral zone of Standoff Rock, where it
formed part of the Bostrychietum together with Bostrychia mixta, Rhizoclonium
ambiguum and Spongomorpha sp. The plant consists of filaments (Figs. 16—18)

1 The publication date of L. W. Dillwyn’s A Synopsis of the British Confervae is cited by many authors

as 1809. In fact it was published in fascicles issued from 1802 to 1809 and for nearly all species the
precise date of publication is printed on the plate.

190 MARINE ALGAE OF GOUGH ISLAND

which measure 30-37 wide with cells whose length is less than to two and a half
times more than their width depending on how recently they have divided. The cell
walls are lamellated and fairly thick. The filaments are unbranched but rhizoids
are present which terminate in slightly digitate haptera (Figs. 16, 17). Cytological
details are not discernible as the cell contents have shrunk. Growth takes place by
intercalary cell division and all stages of this process can be seen in the material
(Figs. 16, 18).

18 Liiiiititits 17

Ou

Fics. 16-18. Rhizoclonium riparium forma validum (Hansg.) Foslie : 16, filament with
rhizoid, m = newly divided cell, y = rhizoid ; 17, filament attached to R. ambiguum, h =
hapteron ; 18, filament with dividing cell.

No previous record of the occurrence of R. ripartum forma validum as a member
of a Bostrychia mixta community has been found but R. riparium has often been
recorded and some of the records probably refer to this forma. In the British
Museum Herbarium there is a specimen of this plant from the spray zone of the
shore at Knysna, South Africa, which has some Bostrychia mixta with it. This
material was collected by Boodle and annotated by E. S. Barton but was not
included by her in her lists of Cape algae. The spray zone is the usual habitat of
this particular Bostrychietum (cf. Post, 1959, p. 498). Also in the British Museum
Herbarium there is material of R. ripariwm forma validum from Kerguelen Island
collected on the ‘‘Erebus”’ and ‘“ Terror’? Expedition and labelled Conferva
tortuosa ® perreptans in J. D. Hooker’s handwriting. Joly’s record (1957, p. 59) of
R. riparium from Brazil probably refers to forma validum since he records the
filaments as being 35—40u wide.

MARINE ALGAE OF GOUGH ISLAND IgI

The name Rhizoclonium riparium forma validum is usually cited as having been
published by Foslie in Wittrock & Nordstedt, Alg. Aq. Dulc. Exsicc. : n. 624 (1884),
but the printed label of this number, which was included in Fasc. 14, reads :

624. Rhizoclonium riparium (Roth) Harv.
Forma valida, crassit. cell. 26-36, altit. 4 minore—24 plo majore.
ayer ar [Habitat].

M. Foslhe.
The second line is clearly a comment on the characteristics of this particular
specimen and not the last part of the name of a new taxon followed by a diagnosis.
This view is confirmed by the list of new taxa in Fasc. 1-20 of the work (Wittrock
& Nordstedt, Alg. Aq. Dulc. Exsicc. 21: 16 (1889)) where the entry under n. 624 has
the same form as the printed label, whereas on the next page there appears :

944. Kh. hieroglyphicum (Ag.) Kiitz. f. calida (Kiitz.) Wittr.

It may well be that, when communicating the specimen to Wittrock and Nordstedt,
Foslie intended that it should be treated as a new forma, for in 1890 he treats the
name Rhizoclonium riparium forma validum as having been published by them on
n. 624 of their set of exsiccata, but this does not alter the fact that they did not do so.

The epithet validum was first validly published for an infraspecific taxon in
Rhizoclonium riparium by Hansgirg in his Prodromus der Algenflora von Bohmen,
the taxon being based on the specimen issued by Wittrock & Nordstedt. Hansgirg
nowhere states the rank to which he assigns the infraspecific taxa which he designates
with roman letters, but it is clear from the way in which his work is set out that
it is higher than variety, designated by Greek letters thus : “ var. 8) ’’, and they
must therefore presumably be treated as subspecies. Hansgirg’s attribution of the
name to Foslie is based on a mistake and its authorship cannot therefore be cited
as “‘ Foslie ex Hansgirg’’; there is no evidence that Foslie ever regarded this taxon
as having a rank higher than forma. De Toni reduced Hansgirg’s R. riparium
b) validum to the rank of a variety in 1889. Although Foslie in 1890 makes no
reference to Hansgirg or De Toni, he cites the same type and his name must be
treated as an alteration of rank.

This taxon is quite clearly delimited. It consists of plants of the Rhizoclonium
type with unbranched filaments 28-45 wide but mostly in the 30-35y range.
The cells are longer than wide unless they have divided very recently and usually
have fairly thick walls, although wall thickness depends to some extent on external
conditions. There has been much confusion as to what the taxon should be called
and in the first place the difference between Conferva tortuosa Dillw. and C. tortuosa
sensu J. G. Agardh (Alg. Medit. Adr.: 12 (1842), non Dillw.) had to be established.
This was done in 1939 by Chapman, who included them in Rhizocloniwm tortuosum
(Dillw.) Kitz. and Chaetomorpha capillaris (Kiitz.) Borg. respectively. He in-
cluded R. riparium forma validum as a synonym of R. tortuosum. Subsequently,
Chapman transferred R. tortuosum to Lola A. & G. Hamel. This genus, as described
by A. & G. Hamel (1929, p. 1094), is distinguished from Rhizoclonium by the posses-
sion of numerous parietal, not few axial, nuclei ; and from Chaetomorpha by having
heterogamous as opposed to isogamous sexual reproduction. The Hamels also said

192 MARINE ALGAE OF GOUGH ISLAND

that Lola was unbranched and only rarely had rhizoids. Chapman seems to have
transferred R. tortuosum to Lola only because it was unbranched as he gives no
details about either its cytology or reproduction. In view of the great variability
of branching throughout the Cladophorales, I do not think this reason enough to
transfer the plant to another genus. In 1931 (p. 25) G. Hamel included in the
genus Lola the species Conferva implexa Harv. and quoted R. tortuosum Kitz. as a
synonym of this. He said that it had 8-18 nuclei but made no comments on the
reproduction. R. tortuosum is quite distinct from C. implexa and I do not think
it is possible to take this statement as evidence of the cytological structure of
R. tortuosum.

In 1955 Koster published a very careful and exhaustive review of the genus
Rhizoclonium in the Netherlands. In this she included R. riparium forma validum.
Whilst agreeing with Chapman that this was the same taxon as R. tortuosum, she
considered that its separation from R. riparium was untenable. In forma validum
she included all plants, with or without rhizoids, with filaments 28-40u wide
and in forma riparium plants, with or without rhizoids, with filaments 17-33
wide. Examination of Dillwyn material (not type) of Conferva tortuosa and Wittrock
& Nordstedt’s n. 624 on which R. tortwosum forma validum is based, both in the
British Museum Herbarium, confirmed that they agree with this concept. The
Dillwyn specimen of C. tortuosa is without rhizoids and has filaments 35-40 wide,
Wittrock & Nordstedt’s material has many rhizoids and the filaments are about
30u wide. I have not been able to make observations on the nuclei of any of the
specimens I have examined as this was not found to be possible on pressed or
formalin-preserved material.

Sinha (1958) published the results of a cytological investigation on this and other
species of the Cladophorales. Included in his results were the following chromosome
counts :

mitosis meiosis
R. riparium (marine) . ‘ ZN 30 yak
R. riparium (brackish and freshwater) : 230
R. riparium forma validum (brackish) 2n = .24
R. riparium forma validum (marine) : ai 24
R. tortuosum (marine) . ; , , Zn — 24
He then wrote: ‘ There is cytological support for Stockmayer’s view that

R. riparium f. validum is a synonym of R. tortuwosum and that R. riparium is a
distinct species whether found in marine, freshwater or brackish situations ”

In view of all the evidence outlined above, I am in no doubt that Conferva tortuosa
and R. riparium forma validum are synonymous and I am also in agreement with
Koster in retaining this forma in the species R. riparium.

CLADOPHORA RADIOSA (Suhr) De Toni, Syll. Alg. 1 : 354 (1889).—Papenf. in Journ.
S. Afr. Bot. 9 : 79 (1943). (Figs. 19-21.)
Conferva vadiosa Suhr in Flora 17 : 741 (1834).—Kiitz., Sp. Alg.: 390 (1849).
Cladophora catenifera Kiitz., loc. cit. (1849) ; Tab. Phyc. 3: 24, t. 83 fig. 1 (1853).—
E. S. Barton in Journ. of Bot. 34 : 193 (1896).—Baards. in Res. Norw. Sci. Exped.
Tristan da Cunha 9 : 12, fig. 2 B (1941).

MARINE ALGAE OF GOUGH ISLAND 193

Window Rock, frequent below low-water neaps on rocks amongst corallines,
18 Feb. 1956, G.J.S.S. 1512. Church Rock, dominant on vertical rock face,

5 Mar. 1956, G.I.S.S. 1529, 1535.
South Africa, Tristan da Cunha, New Zealand, ? Pondicherry, ? Bermuda.

C. radiosa grew very abundantly on Church Rock where it dominated the lower
littoral zone of the rock and extended into the sub-littoral on the sheer southern
face. It was also collected from the sub-littoral zone at Window Rock. Baardseth
recorded it from littoral pools and from the sub-littoral zone on Tristan da Cunha.

Imm

19 20 21

Fics. 19-21. Cladophora radiosa (Suhr) De Toni: 19, terminal branching ; 20, terminal
branching of specimen from Cape of Good Hope labelled C. pellucida by Harvey ; 21,
habit of Gough Island plant.

194 MARINE ALGAE OF GOUGH ISLAND

The plant grows in dense, erect clumps up to 9 cm. high, which consist of numerous
branched fronds. From the base of the fronds downward-growing rhizoids are
produced which interweave and form a mat on the rock. The lowermost cell of
the frond is very large, measuring up to 40 mm. long x 0:45 mm. wide with thick,
lamellated walls up to 70y thick. The plants (Fig. 21) branch from about two-thirds
of the way up ; the main branches are dichotomous and the secondary ones (Fig.
19) are dichotomous or trichotomous. The blunt apical cells measure from 700u.
to I,300u in length and from 200y. to 330 in width, the shorter ones tending to be
the widest. The upper cells measure up to 2,000n long x 200-330u wide and
the walls are about 45u thick. There is a tendency for the cells to be somewhat
swollen at the cross walls and more particularly at the dichotomies. An occasional
swollen, barrel-like cell is present, but there are not enough of these to make them
a characteristic feature of the plant.

The nomenclature and taxonomy of this little-known species of Cladophora have
become somewhat confused. Barton pointed out that Conferva radiosa and
Cladophora catenifera are both the same taxon, and Papenfuss has confirmed this.
Papentuss also attributes the combination Cladophora radiosa to Kiitzing. However,
although Kiitzing in the Specres Algarum included this species under the general
heading of Cladophora, he put it in Species Inquirendae at the end of the genus
and retained the generic name Conferva for it. The first author to call the plant
Cladophora vadiosa was therefore De Toni.

The species is characterized by the very long basal cell, the rhizoidal base and
the dichotomous to trichotomous nature of the branching. It bears some resem-
blance to C. prolifera (Roth) Kiitz. but is much less wiry ; also that species, when
pressed, is green whereas C. vadiosa is brown. The type specimen of C. catenifera
is a specimen from the Cape of Good Hope collected by Harvey and sent to Kiitzing
as C. pellucida. There are two isotypes of C. catenifera in the British Museum
Herbarium (Fig. 20) with which the Gough Island material agrees quite well.
The main difference lies in the rather longer upper cells of the Harvey material.
The relative shortness of the cells of the Gough Island plant may be due to the
extremely severe conditions of thelocality. Baardseth points out that the apparently
barrel-like shape of the cells of C. catenifera as illustrated by Kiitzing was probably
due to the preservation ; it was certainly not a noticeable feature of the Harvey
material in the British Museum Herbarium after it had been soaked and gently
warmed in a weak detergent solution. ©

Chapman (1956, p. 446) recorded this species from New Zealand and pointed out
its similarity to C. feredayi Harv. Chapman gives no cell measurements and
his drawing is not clear enough to enable comparison of the New Zealand material
with material from other localities.

CLADOPHORA RUPESTRIS (L.) Kiitz., Phyc. Gen. : 270 (1843). (Figs. 22-25.)
Conferva rupestris L., Sp. Pl. 2 : 1167 (1753).

Isolda Rock, frequent in mid-littoral rock pools, 6 Mar. 1956, G.I.S.S. 1545.
Europe, North America, Brazil.

MARINE ALGAE OF GOUGH ISLAND 195

Plants of this species grew in abundance attached to the base of Corallina plants
in littoral rock pools on Isolda Rock. The plants (Fig. 22) are up to 5 cm. high and
are fairly stiff; they are composed of long, straight, branching filaments.
Branching (Figs. 23, 24) is mainly dichotomous but sometimes trichotomous,
particularly in the lower parts of the plant ; there is thus no single main axis.
The angle between the branches is always acute and the plants are not spreading.
At the dichotomies, the basal cells of the branches coalesce (Fig. 24), a charac-
teristic of this species. Growth is mainly intercalary. The apical cells are obtuse
(Fig. 24) and measure 200-350 (-500)u long x 60-gou wide. Cells in the

4cm

ZL

l | | | ! 25

100u

Figs. 22-25. Cladophorarupestris (L.) Kiitz. : 22, habit ofplant ; 23, terminal branching;
24, detail of branching and apex, a = apical cell ; 25, sporangia.

196 MARINE ALGAE OF GOUGH ISLAND

upper branches measure 300-700u long x up to 200y wide and cells in the lower-
most part of the plant measure up to 1,050u long x 150u wide. All the cells have
the characteristically very thick, lamellated wall (Fig. 25) up to 45u thick the
structure of which has been investigated by Frei & Preston (1961). The material
was reproducing (Fig. 25), and spores escaping through a pore in the cell wall can be
seen. In this preserved material it is not possible to determine the type of spore.

Mr. C. van den Hoek, who kindly determined the material, said that had it come
from Europe he would have had no hesitation in naming it C. rupestris but that it
might be better to give it this name only tentatively in view of the geographical
isolation of the locality from the main area of distribution. There have been various
records of this species from the Southern Hemisphere, but the only reliable one is
probably that of Joly (1957, p. 57) from Brazil. J. D. Hooker (1845-47, p. 495) and
Dickie (1876, p. 47) recorded the species from Kerguelen, but the material on which
these reports are based (in the Kew and British Museum Herbaria respectively)
is all Sbongomorpha sp. There are several records of C. rupestris from South Africa
but these have also been re-identified as other taxa: for example, Hohenacker,
Meeresalg. 465 and Tyson, Alg. Exsicc. Austro-Afr. 13 have both been identified by
Papenfuss as C. capensis. Hariot (1889, p. 18) recorded the species from Fuegia,
but I was not able to find the specimen on which this record was based in the
Herbarium of the Muséum National d’Histoire Naturelle, Paris. On present
knowledge, therefore, it seems that the distribution of C. rupfestris in the southern
hemisphere is restricted to the Atlantic Ocean.

CODIACEAE!

CODIUM FRAGILE (Suringar) Hariot in Miss. Sci. Cap Horn 5 : 32 (1889).

Anthocodium fragile Suringar in Ann. Mus. Bot. Lugd.-Bat. 3 : 258 (1867) ; Alg. Jap. Mus.
Bot. Lugd.-Bat.: 23, t. 8 (1870) (reimpr. in Natuurk. Verh. Holl. Maatsch. Wet., Ser. 3,
11:235-4..8 (1872)).

Slaughtered Seal Bay, common below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S.
1506. West Point Reefs, frequent in rock pools below spray zone, 6 Mar. 1956,
G.I.S.S. 1540, 1541.

Cosmopolitan.

This species was common in littoral rock pools on Gough Island and was collected
from West Point Reefs and Slaughtered Seal Bay. The material collected consists
of large, robust plants up to 30 cm. tall with branches up to 2 cm. broad. The
branching is irregular ; it is mostly dichotomous but sometimes numerous small
branches are given off over a very small area or sections of branch present a warty
appearance. The utricles are mucronate, and sporangia are present in G.J.S.S. 1506.

Although the species C. fragile is so widespread, Silva has distinguished a number
of subspecies each of which has a fairly restricted distribution. As he has not
finished his work on the species from the South Atlantic, it is not possible to say
which subspecies the Gough Island material represents.

1 Dr. P. C. Silva is at present working on Codium from the South Atlantic and is including the Gough
Island material in his survey.

MARINE ALGAE OF GOUGH ISLAND 197

CoDIUM ADHAERENS Agardh, Sp. Alg. 1 : 457 (1822).—O. C. Schmidt in Bibl. Bot.
gi: 26, figs. 6, 7 (1923).—-Baards. in Res. Norw. Sci. Exped. Tristan da Cunha 9 :
15, fig. 5 D (1941).
Agardhia adhaerens (Agardh) Cabrera in Phys. Sallsk. Lund Arsber. 1823 : 99 (1823).
Church Rock, dominant over considerable areas of shaded vertical rock face,
5 Mar. 1956, G.I.S.S. 1532, 1535.
Distribution not certain, probably chiefly Atlantic.

This species was collected from Church Rock where it was growing on a shaded,
vertical rock face. The specimens agree well with Baardseth’s description of the
plants from Tristan da Cunha.

PHAEOPHAYTA
ECTOCARPACEAE

FELDMANNIA GLOBIFERA (Kiitz.) Hamel in Bot. Notis. 1939 : 67 (1939).—Baards. in
Res. Norw. Sci. Exped. Tristan da Cunha g : 19, fig. 8 c—D (1941).—Kuckuck in
Helgol. Wiss. Meeresuntersuch. 6: 179, figs. 5-10 (1958). (Figs. 26-34.)
Ectocarpus globifer Kiitz., Phyc. Gen.: 289 (1843).—Bornet in Bull. Soc. Bot. France 38 :

358, t. 7 figs. 6, 7 (1891).—Borgesen in K. Danske Vid. Selsk. Biol. Medd. 6 (2) : 48,
figs. 25, 26 (1926).
Slaughtered Seal Bay, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1506.
Europe, Canary Islands, Tristan da Cunha.

The plant was found growing in tufts on Codium fragile in rock pools at Slaughtered
Seal Bay. The tufts are about 2 cm. high and have a rhizoidal base, the filaments
of which weave in among the Codium utricles and anchor the tufts firmly. Kuckuck
described two sorts of tuft in European material and the Gough Island plants re-
semble his northern form in being rather compact and having a dense “ kernel ”’
of branched filaments where most of the reproductive bodies are found surrounded
by an outer fringe of unbranched, lightly pigmented filaments.

The erect filaments are branched, ribbon-like and 40-50u wide. These dimensions
are similar to those given by Baardseth for the Tristan da Cunha material and by
Bergesen for that from the Canary Islands but are rather smaller than in the Euro-
pean specimens described by Kuckuck as being about 70 wide. The meristems
are intercalary except in some young side branches (Fig. 26), which are still entirely
meristematic. The meristems and branches occur in the lower two-thirds of the
tufts and beyond this region the filaments extend into long, scantily pigmented
hairs. Branching is irregular ; many branches arise from the growth of the stalk
cell of old sporangia (Fig. 27). The branches are initially at right-angles to the
filaments which bear them, but gradually the lower wall of the basal cell elongates
more than the upper wall and the angle thus becomes acute.

The plants were reproducing abundantly ; both unilocular and plurilocular
sporangia are present. Both types of sporangium can be seen in a single tuft, but
the two types are not found on the same filament, a condition illustrated by
Kuckuck (loc. cit., figs. 9, 10).

198 MARINE ALGAE OF GOUGH ISLAND

ee ET eal

|

i Es ea LS ed

Fics. 26-34. Feldmannia globifera (Kiitz.) Hamel : 26, meristematic side branch ; 27,
branch growing from basal cell of plurilocular sporangium ; 28, unilocular sporangium ;
29, unilocular sporangium with two-celled stalk ; 30, unilocular sporangium growing in
the remains of an old one ; 31, unilocular sporangium with liberated spores ; 32, pluri-
locular sporangia ; 33, spores being liberated from a plurilocular sporangium ; 34,
germinating spores.

Unilocular sporangia have only been recorded once previously : by Kuckuck
on material from Rovigno (now Rovinj) on the Adriatic coast. In the Gough
Island plants these sporangia (Figs. 28-31) are globose, and mature ones have
a diameter of about 50u. With very few exceptions the sporangia have a one-celled
stalk, but occasional two-celled stalks (Fig. 29) are present. Sometimes a new
sporangium develops inside the ruptured wall of an old one (Fig. 30). Unilocular
sporangia are usually borne laterally on the filaments but occasionally they develop
terminally on young side branches. No unilocular sporangia containing ripe

MARINE ALGAE OF GOUGH ISLAND 199

spores were found, and it was accordingly not possible to ascertain how they
developed and were liberated. These unilocular sporangia differ somewhat from
those described by Kuckuck as being sessile and 70-go0u high x 50-70u wide.
Kuckuck also described a second type of unilocular sporangium. This type was
stalked, globose and about 7oy in diameter, and was thus more like the Gough
Island sporangia ; it differed however in being found characteristically on the same
filament with plurilocular sporangia.

The plurilocular sporangia (Figs. 32, 33) are nearly always stalked and lateral
but occasional sessile or terminal ones are seen. They are solid-looking, ovoid
bodies and mature ones measure up to 80u wide xX I00y long ; this is very similar
in size and shape to those described by Kuckuck, Bornet and others. The spores
differentiate in the sporangium and are liberated by the rupture of the apex of the
sporangium (Fig. 33). Plurilocular sporangia also grow inside the ruptured wall
of old sporangia.

Many liberated spores are present in the material (Fig. 34) ; several had germinated
and others had developed into the few-celled plantlet stage on the surface of the
filaments. It is also common to find spores germinating while still inside the
sporangia. These features were also commented on by Bornet.

The Gough Island plants therefore agree closely with this species as it is known
from other parts of the world. Dr. P. Kornmann, who kindly confirmed the
identification, remarked on this close similarity.

Baardseth’s record of this species from Tristan da Cunha was the first from the
Southern Hemisphere but the species is well known from Europe and the Canary
Islands. Good descriptions of the plant have been given by Bornet, Bgrgesen and
Kuckuck.

GIFFORDIA GRANULOSA (Smith) Hamel in Bot. Notis. 1939 : 67 (1939).—Baards. in
Res. Norw. Sci. Exped. Tristan da Cunha 9g: 21, fig. 8 A (1941).—Kuckuck in
Helgol. Wiss. Meeresuntersuch. 8 : 119, figs. 8,9 (1961). (Figs. 35, 36.)
Conferva granulosa Smith in Sowerby, Engl. Bot. 33 : t. 2351 (1811).

Ectocarpus granulosus (Smith) Agardh, Syst. Alg.: 163 (1824).—Sauvageau in Bull. Stat.
Biol. Arcachon 30: 59, figs. 13, 14 (1933).—Lindauer, Chapman & Aiken in Nova
Hedwigia 3 : 145, t. 82 fig. 4 (1961).

Slaughtered Seal Bay, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1506.
Widely distributed in warm and temperate waters.

A small quantity of this plant was found growing as an epiphyte on Codium
fragile in rock pools at Slaughtered Seal Bay ; it was intermingled with the much
more abundant growth of Feldmannia globifera. The soft, feathery plants are up to
2°5 cm. tall and show the characteristic opposite branching (Fig. 35) of the species
and the cortication of the lower part of the main axis by downward-growing rhizoids
(Fig. 36). The main axis and branches measure 40-70u wide, which is rather
narrow for this species (cf. up to 100u given by Kuckuck) and the material generally
is in rather poor condition. The cells are barrel-shaped and in most cells the
contents have disintegrated ; in some, however, the typical discoid chromato-
phores can be clearly seen. The meristematic regions (Fig. 35) are intercalary

200 MARINE ALGAE OF GOUGH ISLAND

35

Fics. 35, 36. Giffordia granulosa (Smith) Hamel: 35, upper part of plant, m = meri-
stematic region, s = sporangium ; 36, rhizoids growing from lower cells of plant.

and in these parts the cells are wider than long. In the more mature parts, the
cells are more elongated, especially in the branch apices where they are also very
lightly pigmented. The apices are not prolonged into long hairs as they often are
in this species, but are very like those illustrated by Sauvageau.

Sessile sporangia are present on the upper sides of the side branches (Fig. 35).
The young sporangia are quite clearly divided and plurilocular ; although divisions
are not apparent in the mature sporangia, they are almost certainly all plurilocular.
The apparent absence of cell walls is probably due to the generally poor state of
preservation of the material. The sporangia are ovoid and mature ones measure
40-60u. wide x 60-80 long ; this is within the usual size range for this species.

MARINE ALGAE OF GOUGH ISLAND 201

GIFFORDIA MITCHELLIAE (Harv.) Hamel in Bot. Notis. 1939 : 66 (1939).—Baards.
in Res. Norw. Sci. Exped. Tristan da Cunha g : 21, fig. 7 E (1941).—Levring in
Act. Univ. Lund., Ny Foljd, Avd. 2, 56 (10) : 14 (1960). (Figs. 37-41.)
Ectocarpus mitchelliae Harv., Ner. Bor.-amer. 1 : 142, t. 12 fig. G (1851) (reimpr. in Smithson.

Contrib. Knowl. 3 (4) : 142, t. 12 fig. G (1852)) (‘‘ Mitchellae ’’)—Beorgesen in Dansk Bot.
Ark, 2 (2) : 6, figs. 3, 4 (1914) (reimpr. ut Alg. Dan. W. Ind.: 162, figs. 129, 130 (1914)) —
Joly in Univ. Séo Paulo Fac. Fil. Ci. & Let. Bol. 217, Bot. 14 : 72, t. 5 figs. 7, 7a (195°7).

Ectocarpus virescens Thur. ex Sauvageau in Journ. de Bot. 10 : 124, figs. 1-7 (1896).—

Hamel, Phéophyc. France : 29, fig. 7 (1931).

a7

4l

10
Fe eer oF i ear ee

Fics. 37-41. Giffordia mitchelliae (Harv.) Hamel: 37, rhizoidal base of plant ; 38,
branching, m = meristematic region ; 39, detail of cell contents, d = discoid chromato-
phore, m = nucleus ; 40, plurilocular sporangium ; 41, dehisced plurilocular sporangium,

m == meiospore.

202 MARINE ALGAE OF GOUGH ISLAND

Dell Rocks Beach, occasional in sheltered places between boulders and in shallow
pools, 12 Apr. 1956, G.I.S.S. 1558.
Widely distributed in warm and temperate waters.

This species was found growing on rocks in shallow pools and in sheltered places
between boulders at Dell Rocks Beach. The plants are about 5 cm. tall and soft
and feathery like the form from sheltered places on Tristan da Cunha reported by
Baardseth. The base of the plant is rhizoidal (Fig. 37) and the upper part richly
and irregularly branched (Fig. 38) ; the side branches are sometimes secund as
shown in the figure, but this is not a pronounced feature of the plant. The main
axes measure up to 40u. wide with cells up to twice as long as wide. The cells have
discoid chromatophores (Fig. 39). Meristematic regions occur at the base of the
side branches (Fig. 38) ; these branches usually taper somewhat towards the apex
but the apical cells are always obtuse.

Plurilocular sporangia are present (Figs. 40, 41) ; these are usually sessile
and lateral on side branches but are occasionally found on the main axis or terminally
on young side branches. The sporangia are elongated with rather a blunt apex ;
they vary considerably in size, the largest measured being 1204 xX 30p. The
loculi are about 6-7u high x 7—8y wide and thus correspond (like the ones described
by Baardseth from Tristan da Cunha) to the meiospore loculi of Sauvageau (tom.
rol ces UIE

SPHACELARIACEAE

SPHACELARIA FURCIGERA Kiitz., Tab. Phyc. 5 : 27, t. go fig. 2 (1855).—Reinke in
Bibl. Bot. 23: 14, t. 4 figs. 5-13 (1891).—Skottsb. in Wiss. Ergebn. Schwed.
Siidpol.-Exped., Bot. 4 (6) : 58 (1907).—Irvine in Trans. Bot. Soc. Edin. 37 :
28 (1956).—Levring in Act. Univ. Lund., Ny Foljd, Avd. 2, 56 (10) : 16 (1960).
(Fig. 42.)

Fic. 42. Sphacelaria furcigera Kiitz. : branching, propagule on left.

MARINE ALGAE OF GOUGH ISLAND 203

Slaughtered Seal Bay, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1506.
Widely distributed in warm, temperate and colder waters.

This species was found growing as an epiphyte on Codium fragile in rock pools
at Slaughtered Seal Bay. The plant is firmly anchored to the host by means of
rhizoids which weave in among the Codium utricles and form knotted masses at
intervals. Reinke shows the rhizoids of S. furcigera actually penetrating the tissue
of the Sargassum on which it is growing, but the nature of the Codium thallus is such
that the Sphacelaria cannot penetrate its tissues. The plants measure up to 12 mm.
high ; the upper parts are sparingly branched (Fig. 42) and hairless; the
filaments measure 20-40u wide, which accords well with the dimensions given by
Reinke and Irvine ; the plants also bear a close resemblance to the figures of
Kiitzing. Sporangia are absent but occasional bi-radiate propagules are present
(Fig. 42), the appearance of which does not differ markedly from ordinary side
branches.

Some of the material was sent to Dr. Irvine who confirmed the identification and
commented as follows : “I could find no well-developed propagules, though a few
showed clear signs of developing two arms only. The young propagules appear
to be second or even third generation from the base and there is no sign of hairs
either on the propagules or elsewhere. The general dimensions, mode of branching,
number of longitudinal divisions of segments (1-3) and form of propagules seem
right for S. furcigera ; the lack of hairs is probably unimportant as in general I
have found that their presence is very variable, apparently depending on the tem-
perature, at least to some extent, i.e. the higher the temperature the more hairs.
The only point that does worry me a little is the presence here and there of quite
distinct little holdfasts like those of S. pennata ; in general though, the axes spring
from dispersed rhizoids as is usual in this species. There are also very occasional
secondary transverse divisions, but I do not really think that this is of much
significance .”’

Skottsberg recorded the species from Tierra del Fuego and the Falkland Islands.

PUNCTARIACEAE

COLPOMENIA SINUOSA (Roth) Derb. & Solier in Suppl. Compt. Rend. Acad. Sci.
Par. 1: II, t. 22 figs. 18-20 (1856).—De A. Saunders in Proc. Calif. Acad. Sci.,
Ser. 3, Bot. 1 : 164, t. 32 figs. 7, 8 (1898).—Setch & Gardn. in Univ. Calif. Publ.
Bot. 8 : 539, t. 45 figs. 82-86 (1925). (Figs. 43-45; Plate 16 B.)

Ulva sinuosa Roth, Catalect. Bot. 3 : 327, t. 12 (1806).

Midshipman Rock, gelatinous form frequent in lower littoral, 18 Feb. 1956,
GIS5. 1576.

Probably cosmopolitan.

This species was collected only once from Gough Island, where it grew quite
abundantly on rocks of the lower littoral zone of the Midshipman Rock. The plants
form brown, warty, gelatinous cushions which are up to I cm. thick (Plate 16 B).

204 MARINE ALGAE OF.GOUGH ISLAND

In section (Figs. 43-45) the plant body consists of an outer layer, one or
two cells deep, of more or less cubic cells up to about 12y in diameter, which are
usually pigmented. Inside this cortex is the medulla, consisting of colourless,
irregular cells which increase in size towards the centre. The middle of the thallus
is hollow. Pigmented hairs are present ; these grow fairly deeply embedded in
the thallus (Fig. 45) in cryptostomata and are about 7p in diameter. Plurilocular
sporangia were found over much of the surface of the thallus ; the sporangia
(Figs. 43, 44) arise from the outer cells of the cortex and consist of sixteen cells
in two rows of eight. Mature sporangia measure up to 40u high x 8y wide.
Paraphyses are present between some of the sporangia ; they are large and some-
what clavate and measure about 50u X I5yu. The cortex and medulla together are
about 160 thick.

; ; 2208 @)>ys

Nu} i oe, lOul
ee
oy
b F
| i |
L

44

43

Fics. 43-45. Colpomenia sinuosa (Roth) Derb. & Solier: 43, vertical section through
thallus, p = paraphysis, ms = mature sporangium ; 44, vertical section through thallus,
p = paraphysis, ys = young sporangium ; 45, vertical section through thallus, h =
hair, c = cryptostoma.

The type form of C. sinuosa is a balloon-like, hollow ball. Several varieties have
however been described which closely resemble the tuberculate Gough Island plant.
Saunders (tom. cit. : 164) described a new species, C. tuberculata, which was subse-
quently reduced to C. sinuosa forma tuberculata by Setchell & Gardner (tom. cit. :
541). This forma is extremely like the Gough Island plant in external form but
differs in microscopic detail. The cortex is about four cells deep and the sporangia,
which measure up to 25 tall, have cells in a single row, not a double row. The

MARINE ALGAE OF GOUGH ISLAND 205

Gough Island plant also strongly resembles Iyengaria stellata (Berg.) Borg. (1939,
p. 92) externally, although it is smaller than that species (which includes Colpomenta
capensis Levring (1938, p. 20)) in both macroscopic and microscopic detail. The
generic status of [yengaria is rather dubious as it is separated from Colpomenta
only by the absence of paraphyses among the plurilocular sporangia and an allegedly
somewhat different growth process in the thallus. Re-investigation of [yengaria
is needed ; meanwhile the Gough Island plant would seem to differ from it in the
possession of sporangial paraphyses.

In conclusion, I think that this alga comes within the generally accepted variation
range of C. sinwosa, and while it does not externally resemble the type form of this
species, it is yet not sufficiently like any of the described varieties to be referred to
one of them. Tuberculate forms of C. sinuosa have been mentioned by various
authors and of particular interest is that described and illustrated by Joly (1957,
p. 84, t. 4 fig. 3 B) from Brazil, which appears to be very similar, macroscopically,
to the Gough Island plant. Unfortunately, Joly does not give an account of the
microscopic structure and further comparison is thus impossible.

SCYTOSIPHON LOMENTARIA (Lyngb.) Endl., Gen. Pl., Suppl. 3 : 25 (1843).—Setch.
& Gardn. in Univ. Calif. Publ. Bot. 8 : 531, t. 44 figs. 72, 74 (1925).—Baards. in
Res. Norw. Sci. Exped. Tristan da Cunha 9: 31 (1941).—Silva in Univ. Calif.
Publ. Bot. 25 : 258 (1952).

Chorda lomentaria Lyngb., Tent. Hydrophyt. Dan.: 74, t. 18 fig. E (1819).:

Dell Rocks Beach, near low-tide mark on boulder beach, 19 Feb. 1956, G.I.S.S.
I51r ; same locality, frequent in pools and protected channels just above mid-tide
level, 12 Apr. 1956, G.I.S.S. 1559. Midshipman Rock, in standing water in upper
littoral, 18 Feb. 1956, G.J.S.S. 1519. West Point Reefs, in bird-fouled spray-zone
rock pools, 6 Mar. 1956, G.I.S.S. 1536.

Cosmopolitan.

Of the four collections of this plant made on Gough Island, two (r5z9 and 1536)
are of the form which Setchell & Gardner designate as cylindricus minor and the
other two are of that which they call typicus. The plants of the latter come from
the mid and lower shore while the others are from pools, often bird-fouled, on the
upper shore. Baardseth recorded both forms from Tristan da Cunha.

LESSONIACEAE

MACROCYSTIS PYRIFERA (L.) Agardh, Sp. Alg. 1: 47 (1820).—Skottsb. in Wiss.
Ergebn. Schwed. Siidpol.-Exped., Bot. 4 (6) : 80, figs. 91-170, t. 9 (1907).—
Womersley in Univ. Calif. Publ. Bot. 27 : 118, tt. 1, 3, 4 (1954).—G. A. Knox
in Proc. R. Soc. Lond., Ser. B, 152 : 588 (1960). (Figs. 46, 47.)

Fucus pyriferus L., Mant. Pl. Alt.: 311 (1771).

Between The Admiral and The Commodore, forming a belt around the island
between 18 and 55 m., also in gullies, 22 Dec. 1955, G.J.S.S. 1502. Near Penguin
Island, at 55 m., 2 Jan. 1956, G.I.S.S. 1509.

206 MARINE ALGAE OF GOUGH ISLAND

47
Fics. 46, 47. Macrocystis pyrifera (L.) Agardh: 46, holdfast, dh = dichotomous
haptera ; 47, air bladder.

Circum-subantarctic between latitudes 40° S. and 60° S. ; west coast of South
America as far north as Peru ; west coast of North America.

Two collections of M. pyrifera were brought back from Gough Island including
an intact holdfast and an apical frond with the leaflets joined together terminally.
The holdfast (Fig. 46) shows the dichotomous haptera typical of this species. The
pear-shaped air bladders (Fig. 47) at the base of the leaflets are also a characteristic
feature.

Plants of this species formed a band round Gough Island from about 18 to about
55 m. immediately below the Durvillaea band ; it was absent only from the very
exposed part of the western end of the island.

DURVILLAEACEAE

DURVILLAEA ANTARCTICA (Cham.) Hariot in Notarisia 7 : 1432 (1892).—Skottsb. in
Wiss. Ergebn. Schwed. Siidpol.-Exped. 4 (6) : 140, fig. 172 (1907).—Naylor in
Ann. of Bot., New Ser., 13 : 285, figs. 1-8, tt. 6, 7 (1949) ; in Trans. R. Soc.
New Zeal. 80 : 277 (1953).—Lindauer, Chapman & Aiken in Nova Hedwigia 3 :
284, t. 65 (1961).

Fucus antarcticus Cham. in Choris, Voy. Pitt. Monde, Travers. de Cronstadt Chili : 7, t. 7
1822).

ie utilis Bory in D’Urville, Fl. Il. Malouines : 17, 22 (Dec. 18251) (reimpr. in Mém.
Soc. Linn. Par. 4 : 589, 594 (Jan. 18261)), nom. invalid.

1 For these dates see under Jvidaea undulosa, p. 224

MARINE ALGAE OF GOUGH ISLAND 207

Durvillaea utilis Bory in Dict. Cl. Hist. Nat. 9 : 192, t. 49 (Feb. 18261), nom. illegit.; in
Duperrey, Voy. Monde La Coquille, Bot., Crypt. : 65 (Dec. 1827), t. 1 (Jan. 1827), t. 2
fig. 1 (Mar. 1827).—Hook. & Harv. in Hook. f., Bot. Antarct. Voy. I : 176 (1845), 454
(1847).—Hariot in Miss. Sci. Cap Horn 5 : 53 (1889).

Outer Dell Rocks, dominant from mid-tide level to 2 m. below low-tide level on
immobile rocks, occasional in deep rock pools, 18 Feb. 1956, G.I.S.S. 15r0.

Sub-antarctic Islands, South America, New Zealand.

This species dominated the lower littoral and sub-littoral zones of the Gough
Island shores. Two collections were made, one of young plants showing the shield-
like attachment organ and divided, leathery frond ; the other a fragment of a
mature frond showing the spongy, honeycomb texture. The young plants had no
reproductive organs but the mature frond bore female conceptacles. The plants
grew to an immense size with fronds up to 3 m. long and holdfasts at least 7-5 cm.
thick.

D. antarctica is a very characteristic species of the sub-antarctic sub-littoral ; its
distribution is mapped by Knox (1960, p. 589). It does not grow as far north as
Tristan da Cunha in the Atlantic.

In the prefatory part of the Flore des Iles Malouines, D’ Urville gives an account of
the general appearance of an alga and concludes: “‘ M. Chamisso I’avait trés-bien
décrit sous le nom de Fucus antarcticus ; mais M. Bory y a reconnu le type d’un
genre nouveau, et a jugé a propos de l’appeler Durvillaea utilis.’’ In the list of
species, compiled, as far as the cryptogams were concerned, by Bory, the entry
relating to this species reads: ‘27. Durvillaea utilis. Bory. Porro. Le Gentil
(Voyage, tome II, pl. 3.) Fucus antarcticus. Chamisso (Voyage pitt. de Choris.)
Laminaria porroidea. Lamour. (Dict. class. hist. nat. art. Géographie botanique).”’

“ Porra ’”’ of Le Gentil is a vernacular name and the botanical name provided for
the species in his work is a “ pre-Linnaean”’ one: Fucus ramis ex tuberculo rotundo
extentibus, foliis planis, profunde crenatis, pediculatis, pediculis uno versu dispositis
(Le Gentil de la Galaisiére, Voy. Mers Inde 2: 215, t. 3 (1781)). The name
Laminaria porroidea was never validly published, although Lamouroux (Dict. Sci.
Nat. 25 : 189 (1822)) had published Laminaria porra, based on Le Gentil’s “ Porra’’.

In his full account of Durvillaea in the Voyage . . . Coquille, Bory points out that
D’Urville’s account was based primarily on a different alga, Laminaria buccinalis
(L.) Lamour. (= Ecklonia maxima (Osb.) Papenf.), and that Le Gentil’s “‘ Porra ”’
also was not Durvillaea but probably Macrocystis. He states, however, that
Lamouroux had imperfect specimens of Durvillaea from Valparaiso in his herbarium
labelled Laminaria porroidea.

The generic name Durvillaea, and hence the specific name D. utilis, is not validly
published in the Flore des Iles Malouines. D’Urville’s remarks can scarcely be
treated as a description. Even if they could, they would be a description of the
species already described as Fucus antarcticus by Chamisso, and neither they nor
Chamisso’s original description can be regarded as a descriptio generica-specifica, for
the genus, though monotypic, is not founded on a new species.

1 For this date see under Ividaea undulosa, p. 224

208 MARINE ALGAE OF GOUGH ISLAND

When the generic and specific names were validly published by Bory in the
Dictionnaire Classique, he again cited Fucus antarcticus as a synonym of Durvillaea
utilis, the epithet of which is consequently superfluous and illegitimate. In this,
and throughout his works, Bory spelled the generic name Durvillaea, indicating that
he intentionally latinized D’Urville’s name as “ Durvillaeus’’. The “ correction ”’
to ‘‘ Durvillea’’, the form usually adopted, cannot therefore be justified under
Art. 73 of the International Code.

RHODOPHYTA

BANGIACEAE

BANGIA FUSCOPURPUREA (Dillw.) Lyngb., Tent. Hydrophyt. Dan. : 83, t. 24 fig. c
(1819).
Conferva fuscopurpurea Dillw., Synops. Brit. Conferv.: t. 92 (1807).

Standoff Rock, from upper littoral, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1527.
Cosmopolitan.

Only one collection of this species was made and that was from the supra-littoral
zone of Standoff Rock where the Bangia was growing with Porphyra tristanensis,
Hildenbrandia sp. and Verrucaria sp.—a very characteristic supra-littoral
community. Although it was not recorded or collected from any other part of
Gough Island, it is a rather inconspicuous plant when growing and probably occurs
elsewhere on the island.

PORPHYRA TRISTANENSIS Baards. in Res. Norw. Sci. Exped. Tristan da Cunha
9g: 36, figs. 14,15 F (1941). (Figs. 48-51.)

Slaughtered Seal Bay, 22 Dec. 1955, G.I.S.S. 1503. Midshipman Rock, upper
shore, 18 Feb. 1956, G.I.S.S. 1513. Standoff Rock, from upper littoral, 18 Feb.
£056, GI'S.S..7527.

Tristan da Cunha group of islands.

P. tristanensis is the dominant plant of the Gough Island supra-littoral, where it
forms a zone together with a lichen, Verrucaria sp.

Some of the material was sent to Miss J. M. Graves who has studied South African
Porphyra. I am grateful to her for drawing Figs. 48-50 and for the following
description and comparison with P. capensis Kiitz. :

‘‘ The specimens correspond quite well with P. capensis as that species is found in
moderately warm water (e.g. False Bay, Cape). The holdfast has the typically
umbilicate form. In section [Fig. 48] the vegetative cells appear to have one
chromatophore. The cells are not so markedly elongate as are those of typical
P. capensis but I think they fall within the variation range. The protoplasts have
unfortunately shrunken away from the cell walls which makes measurement of the
cell dimensions a little difficult, but as far as I can make out, the vegetative cells are
from 15-25 wide x 50-60 long as seen in transverse section. The ratio length/
breadth therefore varies from 2:1 to 4:1. The majority of the cells measured are
about three times as high as broad as is the case in warm-water P. capensis. A

MARINE ALGAE OF GOUGH ISLAND

AD\Aa (BDO OA «@
D OBESSRHIET |
eee is V

Fics. 48-51.

50

Porphyra tristanensis Baards. :

48, transverse section through thallus,

vegetative cells ; 49, surface view of thallus, c = carpospores ; 50, transverse section
through thallus of G.IJ.S.S. 1513, ¢ = carpospores ; 51, transverse section through thallus
of G.I.S.S. 1503, ¢ = carpospores.

comparison of measurements of P. capensis and the Gough Island plants can be

seen in the following table :

Length of Width of Thickness
vegetative cells in | vegetative cells in of
transverse section | transverse section thallus

Cold-water 100-135 | 12-26. 130-180
P. capensis |

Warm-water 47-68 u. II-19p 70-80
P. capensis

Gough Island 50-60. 15-25 up to gou
Glas 1513

210 MARINE ALGAE OF GOUGH ISLAND

“The plants are all carposporophytic. The number and arrangement of
carpospores in the parent cell is similar to that in warm-water P. capensis. There is
a total of sixteen carpospores per cell and these appear as two rows of four cells
each [Fig. 50] in transverse section. The arrangement in surface view [Fig. 49]
is somewhat more irregular than in P. capensis. The distribution of the fertile
area on the thallus—an irregular marginal zone which is slightly broader at the distal
end than at the sides—corresponds well with P. capensis.

“ To sum up, I would say that the available material corresponds quite well with
warm-water P. capensis.”

I myself thoroughly investigated the plants in G.J.S.S. 1503 and found them to
agree with Miss Graves’s description of G.J.S.S. 1513 as to the vegetative characters.
In this material also, only carposporophytic plants are present and in these plants
there are often up to 32 carpospores per parent cell (Fig. 51) ; this also comes
within the variation range of P. capensis (see Isaac, 1957).

The taxonomy of the genus Porphyra is extremely confused ; with such a simple
structure, the number of characters is small and there seems to be considerable
overlapping between species. Workers on European members of the genus are
gradually elucidating the taxonomy by studying the appearance and development
of living plants and it seems that only by doing this in conjunction with careful
study of type material will it be possible to define specific limits within the genus.
Unfortunately in this case only preserved material is available for study and all
the plants are female. The plants are thus probably dioecious but it may be
that it is a monoecious species in which the spermatia have not yet developed.
I think there is little doubt, however, that this species is the same as that described
by Baardseth from Tristan da Cunha and that this is probably the same as
P. capensis from South Africa.

Levring (1953, p. 464, and 1956, p. 410) published accounts of Porphyra in
Australia and New Zealand and found that, of the two lithophytic species,
P. umbilicalis J. G. Agardh was dioecious with usually 8 carpospores per parent
cell while P. columbina Montagne was monoecious with 32 carpospores per parent
cell. Other workers, however, consider that P. wmbilicalis can be either monoecious
or dioecious and this probably also applies to P. columbina as it does to P. capensis.
It seems, therefore, best to call the Gough Island plants P. tristanensis until the
relationship between this and other southern cold-temperate species has been
elucidated by the study of living material of them all.

GELIDIACEAE

GELIDIUM REGULARE Baards. in Res. Norw. Sci. Exped. Tristan da Cunha g : 52,
figs. 23, 24 A (1941). (Fig. 52.)
Slaughtered Seal Bay, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1507.
Tristan da Cunha (Nightingale Island).
G. regulare was found in littoral rock pools at Slaughtered Seal Bay ; it was
growing with Corallina officinalis, Dermatolithon nodulosum, Lithothamnium
lamellatum and Centroceras clavulatum.

MARINE ALGAE OF GOUGH ISLAND 211

The plants (Fig. 52) measure up to 6 cm. tall. They show the distinct main axis and
abundant, tripinnate branching described by Baardseth. Tetrasporangia are
present. Baardseth pointed out that this species is very similar to the European
species G. latifolium (Grev.) Born. & Thur., but until more is known of the distribution
and variation range of that species it seems best to give the South Atlantic plants a
different name.

5mm Q) ( WA
4, \ 9 y,
fi X NUN HE
Bs NY Sie
= Ave y SE

v4

f

s)
YZ,
RS
Wy ROT'

ae

Fic. 52. Gelidium regulare Baards. : habit of plant, ¢ = tetrasporangia.

CORALLINACEAE

CORALLINA OFFICINALIS L., Syst. Nat., ed. 10, 1 : 805 (1758). (Figs. 53, 54.)
Slaughtered Seal Bay, in pools from water level down to 10 cm., 22 Dec. 1955,
G.I.S.S. 1504 ; same locality, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1507.
Midshipman Rock, dominant in mid-littoral zone on shiny olive-green deposit, 18 Feb.
1956, G.I.S.S. 1515, 1517, 1518. Standoff Rock, top of mid-littoral, 18 Feb. 1956,

212 MARINE ALGAE OF GOUGH ISLAND

G.I.S.S. 1524, 1525, 1526. West Point Reefs, pools below spray zone,6 Mar. 1956,
G.I.S.S: 1541. Isolda Rock, in shallow mid-littoral pools, 6 Mar. 1956, G.I.S.S. 1557;
same locality, upper-littoral pools, 6 Mar. 1956, G.I.S.S. 1553.

All Northern and Southern Hemisphere temperate seas, arctic, antarctic.

C. officinalis was very common on the shores of Gough Island ; it grew in littoral
rock pools, and in two places, Standoff Rock and the Midshipman Rock, it formed
a zone on the rocks at mid-tide level.

The plants (Fig. 53) are small, seldom exceeding 4 cm. in height. They have
a very pronounced main axis bearing densely crowded, slender, pinnate branches
which often branch again. The intergenicula of the main axis and first-order
branches are flattened, the second-order branches are usually thread-like and
terete. Reproductive conceptacles (Fig. 54) are abundant ; they are globose

Imm

Fics. 53, 54. Corallina officinalis L. : 53, habit of plant ; 54, conceptacles (c).

with an apical pore and are occasionally antenniferous. In longitudinal section
the cells of the intergenicula are seen to be in horizontal rows all of much the same
height ; the cells measure about 60-7ou high x 7-gu wide and form lateral
synapses. Each geniculum is composed of a single row of long cells with thick
walls. The plants are frequently covered with epiphytes, Herpostphonia paniculata
being the most common one.

MARINE ALGAE OF GOUGH ISLAND 213

C. officinalis has frequently been recorded from southern cold and temperate
waters, but in only one case (Skottsberg, 1923, p. 67) has the material been described
and this material is obviously different from the Gough Island plants. The Gough
Island plants differ from the type form of C. officinalis in their small size and very
dense branching but I am of the opinion that they come within the variation range
of this species. Dr. Levring, to whom I sent some material, said that it closely
resembled his material from Chile and he thought that it should be included in
C. officinalis (see Levring, 1960, p. 40).

In the collections in the herbaria of the British Museum, Kew and the Muséum
National d’Histoire Naturelle, Paris, there are a few specimens which much resemble
the Gough Island material of C. officinalis and these specimens are variously identi-
fied as C. officinalis, C. cuviert and C. chilensis. The distinctions between these
three and other species, such as C. vancouvertensis, are far from clear and it seems
best to refer the Gough Island plants to C. officinalis.

Corallina goughensis Y. M. Chamberlain, sp. nov. (Figs. 55-60 ; Plate 16 Cc.)
Planta usque ad 4 cm. alta, frondibus numerosis erectis e crusta basali ortis, per
duos longitudinis trientes inferiores simplicibus intergeniculis teretibus, per trientem
superiorem dichotome vel corymbose ramosis intergeniculis compressis. Inter-
geniculorum cellulae dispositae in strata horizontalia extremam partem versus
deorsum curvata ita corticem formantia ; cellulae medianae 35—70y longae, 6-8u
latae, synapsibus lateralibus bene evolutis. Genicula unizonalia ; cellulae usque
ad 250u longae. Conceptacula tetrasporica terminalia, in sinu inter ramulos duos
posita, poris apicalibus. Tetrvasporae zonatim divisae, c. 160u longae, 60y latae.

Isolda Rock, very abundant in lower-littoral rock pools, 6 Mar. 1956, G.I.S.S. 1547
(holotype).

Not known elsewhere.

Plant up to 4 cm. tall with numerous fronds springing from a basal crust. Lower
two-thirds of plant unbranched with terete intergenicula ; upper third branched
dichotomously or corymbosely with flattened intergenicula. Cells of intergenicula
(Fig. 56) in horizontal rows curving out at the sides to form a cortex. Central cells
35-70u long x 6-8yu wide, lateral synapses (Fig. 56) very pronounced. Genicula
unizonal (Figs. 57, 58), cells up to 250u long. Tetrasporic conceptacles (Fig. 59)
present in the angles of the branches, pores apical. Tetraspores (Fig. 60) zonately
divided, about 160y long x 60p wide.

This plant was found growing in littoral rock pools at Isolda Rock where it was
very abundant. Externally, the plant (Plate 16 c; Fig. 55) is characterized
by its dichotomous-corymbose branching, internally by the very striking lateral
synapses between the intergenicular cells (Fig. 56) which appear colourless in
an otherwise stained section.

It was very difficult to decide in which genus this plant should be included. In
the most recent survey of the articulated Corallinaceae, Manza (1937, 1940) divides
the species with only terminal conceptacles into the genera Corallina, Jania,

214 MARINE ALGAE OF GOUGH ISLAND

10u

a a

1Ou,

56

intergenicular
cell

genicular
cell

50u

60

Fics. 55-60. Corallina goughensis Y. M. Chamberlain : 55, habit of plant, ¢ = tetra-
sporic conceptacle ; 56, vertical section of intergenicular cells, s1 = side view of lateral

synapsis, ff = face view of lateral synapsis ; 57, vertical section of geniculus, ge =
genicular cell, eg = extra-genicular portion, ig = intergenicular cell ; 58, detail of extra-
genicular portion of genicular cell (diagrammatic), = pit connexion, c = granular

contents, sg = deeply staining genicular cell wall ; 59, vertical section through frond
g = geniculum, 7 = intergeniculum, ¢ = tetrasporic conceptacle ; 60, tetraspores.

MARINE ALGAE OF GOUGH ISLAND 215

Arthrocardia and Duthiea!. With the exception of Duthiea, which is characterized
by having lateral conceptacle pores, the genera are distinguished by their branching ;
on this basis this plant should be classified as a Jania. However, in the Coral-
linaceae the branching even in a single plant is liable to be extremely variable, and
Suneson (1943, p. 48) has pointed out that there are even forms of Corallina officinalis
which are dichotomous. Most authors distinguish Corallina from Jania by the
length of the genicular cells, which are at least twice as long as the intergenicular
cells in Corallina, and are about the same length as the intergenicular cells in Janda.
On this basis I have classified this plant as a Corallina. The material is somewhat
similar to Jania digitata Manza but the intergenicula are more flattened in
C. goughensis and the upper branches are less regularly dichotomous than those
of J. digitata.

Dermatolithon nodulosum Y.M. Chamberlain, sp. nov. (Figs. 61-65 ; Plate 17.)

Planta horizontaliter crustaceo-expansa, in statu vivo rosea, ad 2 cm. crassa,
nodulos calcarios permultos confertim ferens. Crusta basalis ad 700w crassa,
plerumque in vaginam circa nodulos producta, constata e strato supremo simplici
cellulis epithallinis parvis planis, stratis medianis pluris horizontalibus cellulis
perithallinis ad 50u longis et roy latis, strato infimo simplici cellulis hypothallinis
obliquis ad 80y longis et Io latis. Noduli e perithallo bene evoluto cellulis ad
100 longis et 15y latis constati. Cellulae perithallinae et hypothallinae omnes
synapsibus secundariis lateralibus apicem cellulae versus munitae. Conceptacula
tetrasporangiifera in perithallo praesentia, plerumque supra superficiem crustae
eminentia, desuper visa circularia, diametro interno ad 250u metienti, ad 150y alta.
Tetrasporae zonatim divisae.

Slaughtered Seal Bay, in pools from water level down to 10 cm., 22 Dec. 1955,
G.I.S.S. 1504 ; same locality, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1507.
Midshipman Rock, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1514. West Point Reefs,
pools below spray zone, 6 Mar. 1956, G.I.S.S. 154r ; same locality, margins of pools
below spray zone, 6 Mar. 1956, G.I.S.S. 1543. Isolda Rock, very abundant in
shallow mid-littoral pools, 6 Mar. 1956, G.I.S.S. 1550, 1551 ; same locality, upper-
littoral pools, 6 Mar. 1956, G.I.S.S. 1553 (holotype).

Not known elsewhere.

Plant (Plate 17 A) pink when living, up to 2 cm. thick, bearing nodules (Plate 17 B ;
Fig. 61). Basal crust (Fig. 63) up to 700 thick, composed of a single layer
of small flat epithallial cells, horizontal layers of perithallial cells measuring up to
50u long X roy wide and a single layer of oblique hypothallial cells measuring up to
80 long xX Iou wide. Nodules (Figs. 61, 62, 64) composed of a much-developed
perithallus with cells roou long x I5u wide. All perithallial and hypothallial cells
have secondary, lateral synapses which occur towards the top of the cell. Basal
crust often extending to form a sleeve round the nodules (Fig. 62). Tetrasporic
conceptacles (Figs. 62, 65) present in perithallus, usually projecting a little
above the surface. Conceptacles circular in surface view, in section having a
diameter up to 250u and being up to 150u. deep. Tetrasporangia zonately divided.

} This name, Duthiea Manza (1937), is a later homonym of Duthiea Hack. (1895) (Gramineae).

Fics.
nodule showing direction of cell rows ; 62, vertical section through nodule showing
position of conceptacles and sheath of basal crust cells (b), c = conceptacle ; 63, details
of cells of basal crust in vertical section, e = epithallus, p = perithallus, h = hypothallus ;
64, detail of nodular cells in vertical and transverse section, sf = face view of secondary
synapsis, ss = side view of secondary synapsis, p = primary pit connexion ; 65, trans-
verse section of tetrasporic conceptacle.

MARINE ALGAE OF GOUGH ISLAND

lOul
|imm

level B

level C

level B

4p , i

rt
CT) Sip.

65

Dermatolithon nodulosum Y.M. Chamberlain : 61, vertical section through

D. nodulosum was very abundant on the shores of Gough Island ; its conspicuous
pink crusts grew on the margins of deep rock pools and over the rock surface of
shallow pools ; occasionally it grew on the open rocks.

The size and habit of this plant are unusual for a Dermatolithon, most species of
which have a very thin thallus and are epiphytic or epizoic. Two other species,
D. hapalidioides (Crouan) Foslie and D. papillosum (Zanard.) Foslie, have been
recorded as growing on rocks in temperate waters although both are more commonly
epiphytic or epizoic. D. nodulosum is similar to these species in microscopic.

MARINE ALGAE OF GOUGH ISLAND 217

structure but is a very much larger plant and has coarser nodules. The other two
species have so far only been recorded from the European and North African coasts
(Hamel & Lemoine, 1952, pp. 61, 63).

In young plants, only the basal crust is present. The epithallial and outer
perithallial cells (Fig. 63) of this crust have homogeneous, rather dense contents,
the inner perithallial cells become vacuolated and the hypothallial cells are also
vacuolated. The nodules are composed almost entirely of horizontal rows of large
perithallial cells (Figs. 61, 64), which are almost devoid of contents and form
numerous secondary lateral synapses with the cells round them. What little remains
of the cell contents usually collects round the primary pit connexions and secondary
synapses.

Only tetrasporangial conceptacles were found. These have one central pore
(Plate 17 c; Fig. 65). The tetraspores are usually divided zonately into four
but bispores are also present. The conceptacles do not cave in after the spores have
been shed and many empty ones can be seen buried in the older parts of the
perithallus (Plate 17 c).

LITHOTHAMNIUM NEGLECTUM (Foslie) Foslie in Deutsch. Stidpol.-Exped. 1901-03
8 : 207 (1908) ; Contrib. Monogr. Lithothamnia, ed. Printz : 43, t. 9 figs. 1-4
(1929).—Lemoine in zme Exped. Antarct. Frang. 1908-10, Melobesiées : 14
(1913) ; apud Cotton in Journ. Linn. Soc. Lond., Bot. 43 : 196, t. 9 fig. 2, t. 10
figs. 3-6 (1915) ; in K. Svensk. Vetenskapsakad. Handl., Ny Foljd, 61 (4) ; 12
(1920)... (Big: 60:5 Plate 28.)

Lithothamnium muelleri forma neglectum Foslie in K. Norske Vidensk. Selsk. Skr. 1899 (5) :
17 (1900).

Slaughtered Seal Bay, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1507.
Isolda Rock, frequent in lower-littoral rock pools, 6 Mar. 1956, G.I.S.S. 1548 ; same
locality, upper-littoral pools, 6 Mar. 1956, G.I.S.S. 1553.

Antarctic and sub-antarctic.

Two collections (G.I.S.S. 1507, 1548) of this species come from rock pools at
Slaughtered Seal Bay and Isolda Rock respectively. These plants are rather
similar both macroscopically and microscopically. They have leaf-like pink thalli
(Plate 18 A) with concentric markings. The thallus (Fig. 66) is up to about
400p. thick ; it has a single layer of epithallial cells, below which is the perithallus.
The perithallial cells run vertically below the epithallus and measure up to 10u wide
x 17p long ; they run horizontally in the middle of the thallus and curve downwards
below to the hypothallus, the cells of which measure up to 50u long xX 7u wide.
Tetrasporic conceptacles (Plate 18 c) occur in the perithallus ; these measure up to
300% wide xX 200u deep and open through numerous pores. The tetraspores are
zonately divided and measure up to 145y long x 70 wide.

The third collection (G.I.S.S. 1553) comes from Isolda Rock where, together with
Lithophyllum sp., the plant formed a conspicuous band on the rocks in the upper
mid-littoral zone. This plant (Plate 18 B) was white when growing, probably due to
bleaching in its exposed position. A similar band was reported to occur in other

218 MARINE ALGAE OF GOUGH ISLAND

10u

Fie I ae coe sa ep ee nF Sp (0 |

66

Fic. 66. Lithothamnium neglectum (Foslie) Foslie : vertical section through thallus of
G.I.S.S. 1507, h = hypothallus, p = perithallus, e = epithallus.

parts of the island but no other collection from it was made. This plant is smaller
than the preceding ones both microscopically and macroscopically and it probably
represents Foslie’s L. neglectum forma fragile (1908, p. 208).

The two forms are well illustrated in Foslie’s photographs (1929, t. 9) of the
species, G.I.S.S. 1553 resembling Foslie’s figs. 1-3 and G.J.S.S. 1507 and 1548 his

fig. 4.

GIGARTINACEAE

IRIDAEA LAMINARIOIDES Bory in Duperrey, Voy. Monde La Coquille, Bot., Crypt. :
105 (18283), t. 11 fig. 1 A—-D (18271).—Kititz., Sp. Alg. : 726 (1849) ; Tab. Phyc.
17: 3, t.8 figs. c, d (1867).—Montagne in C. Gay, Hist. Fis. & Polit. Chile, Bot.
8 : 352 (1854).—Laing in Chilton, Subantarct. Is. New Zeal. 2 : 506 (1909).—
Levring in Act. Univ. Lund., Ny Féljd, Avd. 2, 56 (10) : 56(1960). (Figs. 67-71.)
Ividaea heterococca Kiitz., Tab. Phyc. 17 : 4, t. 11 (1867).
Iridaea cordata sensu Cotton in Journ. Linn. Soc. Lond., Bot. 43 : 176 (1915) pro parte ;

non J. G. Agardh.

Ivridophycus laminarioides (Bory) Setch. & Gardn. in Proc. Nation. Acad. Sci. Wash. 22 :
470 (1936) ; in Univ. Calif. Publ. Bot. 19 : 206 (1937).

1 For the dates of the natural history parts of the Voyage ... Coquille see Sherborn & Woodward in
Ann. & Mag. Nat. Hist., Ser. 7, 7 : 392 (1901).

MARINE ALGAE OF GOUGH ISLAND 219

Ividophycus boryanum Setch. & Gardn. in Proc. Nation. Acad. Sci. Wash. 22 : 470 (1936),
nom. nud.

Ividophycus boryanum Setch. & Gardn. in Univ. Calif. Publ. Bot. 19 : 202 (1937).

Iridaea boryana (Setch. & Gardn.) Skottsb. in K. Svensk. Vetenskapsakad. Handl., Ser. 3,
19 (4) : 82 (1941).—Levring in Ark. fér Bot. 31a (8) : 13 (1944) ; in Act. Univ. Lund.,
Ny Foljd, Avd. 2, 56 (10) : 55 (1960).

Midshipman Rock, Porphyra zone, 18 Feb. 1956, G.I.S.S. 1514. Standoff Rock,
dominant on less-exposed part of the rock above the Corallina zone, 18 Feb. 1956,
G.I.S.S. 1526. Isolda Rock, occasional in upper littoral on ridges between pools,
6 Mar. 1956, G.I.S.S. 1552.

Widespread in southern South America, Falkland Islands, Kerguelen, Crozet
Islands, Auckland Island, ? North America.

I. laminarioides was one of the most common littoral algae on Gough Island.
On moderately sheltered boulder beaches it formed a conspicuous mid-littoral zone
of bright orange-brown which was bounded above by a Blidingia zone and below
often merged into a Rhodoglossum zone. It has been recorded as a dominant littoral
plant in other regions, such as Chile (Levring, 1960) and Kerguelen and the Crozet
Islands (Levring, 1944).

The Gough Island plants (Figs. 67, 68) grow from communal, fairly thin basal
crusts which spread over the rocks, each crust giving rise to numerous erect fronds.
According to Dr. Holdgate, the smaller, orange-brown plants grew in the more
exposed places while larger plants, which were usually more violet in colour, came
from the sheltered channels and crannies between the boulders. The material
collected measures up to 20 cm. long but is mostly under 9 cm. ; plants from more
sheltered places undoubtedly grew longer than this, however. All the plants have
a small, terete stipe above the basal crust; above the stipe is a strongly channelled
apophysis from which develops the smooth-edged blade which typically has a
cuneate base. Only one small piece of the material was found to be fertile ; this
(Fig. 68) has tetrasporangia along one side of the edge of the blade. In transverse
section (Fig. 69) the blade can be seen to consist of a central medulla of widely
spaced, very thin and elongated cells ; towards the edge the cells become shorter
and fatter and interweave more noticeably ; finally filaments of up to five,
small, oblong cells are produced which lie at right-angles to the surface of the thallus
and form a densely packed cortex. The tetrasporangia develop in the medulla ; the
intercalary development of the tetrads of spores can be seen when they are young
(Fig. 70) and ripe tetrads measure up to about 35y in diameter (Fig. 69).

Various species of Ividaea occur in great abundance on rocky sub-antarctic
shores. The fact that small samples of Jvidaea from this area have frequently been
collected and often identified without reference to previous collections has resulted
in a confused picture of the taxonomy of the genus. Setchell & Gardner in their
1937 paper made the first real attempt since Bory’s account in the Voyage...
Coquille to sort out the taxonomy of the South American species, and their con-
clusions have been accepted by most subsequent authors. Setchell & Gardner
also changed the name of the genus to Ividophycus since Iridaea Bory is a later

220 MARINE ALGAE OF GOUGH ISLAND

10u

67 68

Fics. 67-71. Ividaea laminarioides Bory : 67, habit of plant ; 68, plant with tetra-
sporangia, b = blade, ¢ = tetrasporangia, a = apophysis, s = stipe, d = basal disk ; 69,
transverse section through blade with mature tetraspores, c = cortex, m = medulla,

t = tetraspore ; 70, transverse section of blade with young tetraspores, m = medulla,

t = tetraspore ; 71, transverse section of blade of Bory’s type specimen, c = cortex,
m = medulla.

homonym. Iridophycus has not been generally accepted, however, and Iridaea
Bory has been conserved. There is an error in the entry referring to it in the list of
nomina generica conservanda (Internation. Code Bot. Nomencl. Montreal 1959 : 205
(1961)) ; this reads: “‘ Ividaea Bory, Dict. Class. Hist. Nat.9g: 15. 1826”. On

MARINE ALGAE OF GOUGH ISLAND 221

the page cited Bory used the spelling Jvzdea, on the next page Iridea twice and
Iridaea twice, and on p. 192 of the same volume Jvidaea ; in his other publications
Bory always used the spelling Ividaea.

Among the species of Jvidaea described by Bory is I. laminartoides, the type
material of which was collected at Concepcion, Chile, by D’Urville during the voyage
of ‘‘ La Coquille’. Setchell & Gardner (1937, pp. 202, 206) considered that the type
material—the smaller and larger plants of which Bory thought to be juvenile and
adult stages of the same species—represented two separate species. They called the
adult plants (Bory, 1827, t. 11 fig. 1 D, E) Iridophycus laminariordes and the so-called
juvenile plants (fig. 1 A, B, c) I. boryanum. This last species they considered to be
the same as Kiitzing’s Ividaea heterococca but did not give it that name because in
their opinion the name was based on a monstrosity, the type material carrying a
blue-green epiphyte mistaken by Kiitzing for reproductive organs of the Ividaea.
Various authors (e.g. Skottsberg and Levring in his 1960 paper) have used Setchel]
& Gardner’s classification of the genus but both Skottsberg and Levring pointed
out that the separation of the two species is unsatisfactory since many intermediate
forms occur. I had the opportunity of examining Bory’s type material of
I. laminariovdes in the herbarium of the Muséum National d’ Histoire Naturelle, Paris,
and I have also studied much other herbarium material of Jvidaea in the Paris,
British Museum and Kew Herbaria. In my opinion, Bory’s type material does
represent two species but I think that the plants on which Bory’s fig. I A, B, Cc and D
were based are all the same species, D perhaps being from a more sheltered position
on the shore, and that E (which has a ciliate margin) is a different species. As the
fragment of E drawn by Bory represents all the material there is, I do not think it is
possible to decide what species it is. J. laminariotdes is a very variable species, but
whilst the blade and apophysis may sometimes have irregular outgrowths, they are
never regularly ciliate like Bory’s fig. 1 E. The small form (Bory’s A, B and c), with
which most of the Gough Island material agrees very closely, usually dries to a shade
of brown if pressed when fresh while the larger form (D) is often more red or purple.
As far as I could see when I examined the type material, the lower left-hand blade
of D is in fact a separate plant and was never part of the main frond as shown by
Bory ; the angle it makes with the main frond looks most unnatural in the drawing
too.

The distinctive features of J. laminarioides are the very short stipe, the
pronounced apophysis which is channelled and has a smooth edge, and a blade with
a cuneate base and a smooth edge ; the blade may or may not be divided. I also
think that the fronds usually arise from a communal basal crust ; certainly those in
the Gough Island material did so and much of the herbarium material I examined
appeared to show this too ; unfortunately it was usually impossible to make sure
about this since dried material of Jvidaea disintegrates very quickly when soaked out.
This feature of the communal basal crust has been used by Setchell & Gardner
(1937, p. 211) as one of the diagnostic characters of another of their new species,
Inidophycus caespitipes. Skottsberg (1941, p. 82) commented on and published a
drawing of this species and said how similar the erect fronds are to those of Ividaea

222 MARINE ALGAE OF GOUGH ISLAND

boryana. Iridophycus caespitipes is probably only a form of Ividaea laminarioides
and the more pronounced development of the basal crust could well be a response
to the very exposed habitats from which it has been collected.

While working on the Ividaea laminarioides group of species, I made sections of
the blade of a considerable number of herbarium specimens. I found a wide range of
variation in the tissues, the cells in some being much more densely packed than they
were in others. In the filaments of the cortex, there was a variation both in the
number of cells in the filaments and in the size and shape of the cells. Some of the
variation may have been due to the method of pressing the specimens and I could not
see that there was any significant correlation between the detailed microscopic
structure of the blade and the external appearance of the plant. The observations of
Setchell & Gardner as to the microscopic characteristics of their various species
seem to have been based on a rather small sample in each case. The appearance of
the cross-section of a blade from Bory’s type specimen of J. laminarioides (Fig. 71)
was rather different in the cortical region from the Gough Island material (Fig. 69)
but I do not think the difference is great enough to warrant specific separation of
the two plants. The only plant of Ividophycus caespitipes that I have been able to
examine is the fragment in the Kew Herbarium collected in the Falkland Islands by
Mrs. Vallentin and sent to Setchell & Gardner by Cotton as Ividaea cordata (Setchell
& Gardner, 1937, p. 212). The specimen is so incomplete that it gives no idea of the
external characteristics of the species, but in cross-section the cortical cells of the
blade are very considerably larger than those of all the other material which I
sectioned. For this reason I have not included Ividophycus caespitipes as a synonym
of Ividaea laminarioides.

It is quite clear that, before the taxonomy of Ividaea can be satisfactorily worked
out, much more work will have to be done with fresh material and field observations
made by phycologists. Until this has been done, I think it is difficult to be sure
whether the North American plants which are called I. laminarioides really belong to
the same taxon as the South American ones, and also to sort out the relationship
between J. laminarioides and other very similar species such as J. micans Bory and
I. obovata Kiitz.

IRIDAEA UNDULOSA Bory in D’Urville, Fl. Il. Malouines: 22 (Dec. 1825)! ; in
Mém. Soc. Linn. Par. 4 : 594 (Jan. 1826).—Skottsb. in K. Svensk. Vetenskaps-
akad. Handl., Ser. 3, 19 (4) : 85, figs. 4-7 (1941).—Levring in Act. Univ. Lund.,
Ny Foljd, Avd. 2, 56 (10) : 57 (1960). (Figs. 72, 73.)

Ividaea crispata Bory in Dict. Cl. Hist. Nat. 9 : 16 (Feb. 1826).—Levring, loc. cit. (1960).

Ividaea augustinae Bory in Duperrey, Voy. Monde La Coquille, Bot., Crypt.: 108 (1828),
t. 12 (1827), nom. illegit.—Kiitz., Sp. Alg.: 726 (1849).—Montagne in C. Gay, Hist. Fis. &
Polit. Chile, Bot. 8 : 353 (1854).

Ividaea micans var. ciliolata Hook. & Harv. in Hook., Lond. Journ. Bot. 4 : 263 (1845).
Ividaea cordata var. ciliolata (Hook. & Harv.) Hook. & Harv. in Hook. f., Bot. Antarct.
Voy. I : 485 (1847).
Ividaea ciliata Kiitz., Sp. Alg.: 726 (1849) ; Tab. Phyc. 17 : 4, t. 10 (1867).—Baards. in

1T am indebted to M. R. Deélepine for a photocopy of this paper.

MARINE ALGAE OF GOUGH ISLAND 223

Res. Norw. Sci. Exped. Tristan da Cunha 9 : 76, fig. 41 E (1941).—Levring, loc. cit. (1960).

Iridophycus undulosum (Bory) Setch. & Gardn. in Proc. Nation. Acad. Sci. Wash. 22 : 471

(1936) ; in Univ. Calif. Publ. Bot. 19 : 217, tt. 25, 26 (1937).—W. R. Taylor in Pap.
Mich. Acad. Sci. Arts & Lett. 24 : 146 (1939).

Iridophycus crispatum (Bory) Setch. & Gardn. in Proc. Nation. Acad. Sci. Wash. 22 : 471
(1936) ; in Univ. Calif. Publ. Bot. 19 : 221 (1937).

Iridophycus ciliatum (Kiitz.) Setch. & Gardn. in Proc. Nation. Acad. Sci. Wash. 22 : 471
(1936) ; in Univ. Calif. Publ. Bot. 19 : 220 (1937).

Slaughtered Seal Bay, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1505.
Church Rock, abundant on vertical rock face, 5 Mar. 1956, G.I.S.S. 1528, 1535.
Isolda Rock, frequent in mid-littoral rock pools, 6 Mar. 1956, G.I.S.S. 1546. Window
Pool, in sheltered shallow pool, 12 Apr. 1956, G.I.S.S. 1556.

Southern South America, Falkland Islands, Tristan da Cunha.

Iridaea undulosa occurred quite frequently on the shores of Gough Island ; it was
most often found in littoral rock pools but was also collected from the Cladophora
vadiosa zone just above and below mid-tide level on the sheer southern face of

og 9868
7 PRP

i A

Fics. 72,73. Ividaea undulosa Bory : 72, habit of plant, a = apophysis, e = emergences,

b = blade, ¢ = tetrasporangia ; 73, transverse section of blade, c = cortex, m = medulla,
t = tetrad of spores.

224 MARINE ALGAE OF GOUGH ISLAND

Church Rock. The plants collected measure up to 17 cm. tall x 12 cm. wide
(Fig. 72), are more or less ovate in shape and are of a deep, purple-red colour and
a rather leathery texture when alive. The apophysis is flat or very slightly
channelled and always bears obvious emergences ; it is unbranched but occasionally
the emergences are large enough to look like young plants. The base of the blade is
cuneate to reniform and the margin is, in most cases, ciliate ; in some younger plants
itis smooth. Many of the plants are tetrasporic but no sexual reproductive organs
were found. In section (Fig. 73) the blade can be seen to have the usual Ividaea
structure ; the long, thin cells of the medulla are widely scattered and the cortex
consists of three-celled filaments of fairly large cells. The very wide spacing of
the cells may be more apparent than real ; the plants are extremely gelatinous and
when sectioned the gelatinous substance immediately swells so much that it pushes
the cells apart. Tetrads of spores develop in sporangia in the medulla in an inter-
calary manner characteristic of this genus.

Setchell & Gardner (1937, pp. 215-222) divided the South American species of
Invidaea with ciliate margins and unbranched apophyses into three species, the
specific delimitations depending upon whether the base of the blade was cuneate to
reniform or cordate. Skottsberg (1941, pp. 85-88) considered that these inter-
specific boundaries were untenable and that the three species should be treated as
forming one taxon, a view which I share. The distribution seems to be limited to
the Atlantic and Pacific coasts of South America and some South Atlantic Islands.

The question of whether the name Iridaea undulosa was validly published in
D’Urville’s Flore des Iles Malouines is not easy to decide. That work as a separate
publication certainly antedates the ninth volume of the Dictionnaire Classique
d’ Histoire Naturelle, and so also probably does the sixth part of the fourth volume of
the Mémoires de la Société Linnéenne de Paris, in which it was also printed.!

In the Flore des Iles Malouines there is a general but not a botanical description of
Iridaea micans in the prefatory part by D’Urville, and in the list of species, for the
cryptogamic part of which Bory was responsible, there is :

25. Iridaea micans. Ovato reniformis, undulosa, tenerrima, basi subcordata.

Cap. (fide Gaud.).
26. a. I. undulosa. Laevis, fronde ovato-conica, basi obconica, crassiuscula,
undulato-crispa.
b. I. papillosa.
Nowhere in this work is there an indication that Iv:daea is a new genus, unless this
can be read into the statement : “‘L’éclat de ses couleurs lui a valu le nom d’Jridea
micans”’.

1 Flore des Iles Malouines. Title-page dated 1825 ; not noticed in the Bibl. France ; received at
weekly meeting of Paris Academy of Sciences on 9 January 1826 (Proc.-Verb. Acad. Sci. Par. 8 : 332
(1918)) ; therefore presumably December 1825.

Mém. Soc. Linn. Par. 4 (6). Cover dated Jan. 1826; not noticed in Bibl. France but notices of
other numbers suggest that it ran close to schedule ; not received by Paris Academy of Sciences ;
date on cover (Jan. 1826) probably correct.

Dict. Cl. Hist. Nat.9. Title-page dated Feb. 1826 ; noticed in Bibl. France 15 : 154 (25 Feb. 1826) ;
received at weekly meeting of Paris Academy of Sciences on 20 March 1826 (Proc.-Verb. Acad. Sci.
Par. 8 : 359 (1918)) ; therefore February 1826.

MARINE ALGAE OF GOUGH ISLAND 225

,

However, Bory’s article headed “ Iridée, Ividea’”’ in the ninth volume of the
Dictionnaire Classique, which appeared only a few weeks later, begins “‘ Le genre
ainsi designé par Stackhouse, ne pouvant étre conservé selon Lamouroux, nous
adoptons ce nom pour un genre nouveau de Fucacées’’. The species he describes in
this article include J. micans but not I. undulosa. In his fuller treatment of the
genus in the Voyage . . . Coquille, he treats I. undulosa, and I. crispata, described in
the Dictionnaire Classique, as synonyms of the new and _ superfluous name
I. augustinae.

Unless Ividaea micans and I. undulosa are regarded as having been published in the
Flore des Iles Malouines as species of the genus Ividea Stackh., they are not validly
published there. The spelling of the generic name in the part of that work for which
Bory was responsible differs from Stackhouse’s, and we know from Bory’s subsequent
publications that a different genus was intended. However, in deciding whether a
name is validly published in a particular publication, matter published later cannot
be taken into account. The decision on the place of valid publication of these two
names therefore turns on the answer to the question : had neither vol. 9 of the
Dictionnaire Classique nor the Voyage... Coquille been published, would J7vidaea in
the Flore des Iles Malouines have been considered as a mis-spelling of Ividea Stackh.
or as the name of a new genus for which no definition was provided? The definitions
of the species in the Flore des Iles Malouines are inconsistent with membership of a
genus defined thus : “ Jvidea. Fronde cartilaginea, tereti ; ramosissima, pinnata :
ramis oppositis, supra decompositis, capillaribus . . . Fructificatio incognita .”
(Stackhouse, Ner. Brit., ed. 2 : ix (1816)). They would nevertheless at the time
have been regarded as belonging to the same group of algae as Ividea Stackh., and
their names must therefore be regarded as having been validly published as names of
new species of that genus. Their subsequent transfer to Ividaea Bory cannot be
regarded as the making of new combinations, and hence there is the anomalous
situation that these two correct names were validly published earlier than was the
name of the genus which forms their first part.

RHODOGLOSSUM REVOLUTUM Baards. in Res. Norw. Sci. Exped. Tristan da Cunha :
78, figs. 39 C, 40 E, 41 D (1941). (Figs. 74-77.)

Midshipman Rock, Porphyra zone, 18 Feb. 1956, G.I.S.S. 15174. Standoff Rock,
occasional in mid-littoral coralline zone, 18 Feb. 1956, G.IJ.S.S. 1524. Church Rock,
occasional on vertical rock face, 5 Mar. 1956, G.I.S.S. 1534. West Point Reefs,
pools below spray zone, 6 Mar. 1956, G.I.S.S. 1541. Isolda Rock, in upper littoral
on ridges between pools, 6 Mar. 1956, G.I.S.S. 1552 ; same locality, upper-littoral
pools, 6 Mar. 1956, G.I.S.S. 1553. Window Pool, abundant in sheltered shallow
pool, 12 Apr. 1956, G.I.S.S. 1556.

Tristan da Cunha (Inaccessible Island).

R. revolutum was a common plant of the Gough Island littoral. It grew on rocks
and in pools in the mid-littoral and in some cases, particularly on rather less exposed
beaches such as Dell Rocks Beach, it formed a zone below the Ividaea laminarioides
zone. The only previous record of this species is its original gathering from the

226 MARINE ALGAE OF GOUGH ISLAND

littoral zone of Inaccessible Island. Baardseth gives no indication of its abundance
there, but as the species is not mentioned in the ecological section of Baardseth’s
work it can be assumed that it was not present in any great quantity and certainly
did not dominate a zone as it did in places on Gough Island. It would seem
therefore that this is a plant of very restricted distribution which attains its
maximum development on Gough Island.

The plants (Fig. 74) consist of a spreading basal crust from which arise erect
fronds in all stages of development. The fronds are fleshy and of a dark reddish-
brown colour ; the lower half of most plants is entire ; above this the plants
usually branch dichotomously up to four or five times but occasional unbranched

10u

Fics. 74-77. Rhodoglossum revolutum Baards.: 74, habit of plant ; 75, transverse
section through blade, m = medulla, c = cortex ; 76, transverse section through blade
with tetraspore mother cells (¢mc), c = cortex ; 77, mature tetrad of spores.

plants are present. The margins of the fronds are entire and frequently inrolled.
The fronds measure up to 6 or 7 cm. high and are extremely gelatinous, which
causes them to shrink excessively when pressed. Baardseth describes the plants
as being 2 or 3 cm. high, but this measurement must have been based on dried
material ; the isotypes in the British Museum Herbarium measured up to 3 cm.
on the herbarium sheet, but one specimen which was 2 cm. high when dry expanded
to 3-5 cm. when soaked in water. The Gough Island plants measure up to I cm.
across but are usually about 5 mm.; Baardseth recorded 4-8 mm. for the width.
In section the fronds are flat except at the very bottom where they are terete.
The frond is composed of a cortical region of filaments of small elongate cells at

MARINE ALGAE OF GOUGH ISLAND 227

right-angles to the surface of the thallus and an inner medulla of long, thin branching
cells which anastomose with one another (Fig. 75).

Female and asexual reproductive organs are present. The carposporophytes
develop on the concave surface of the upper parts of the thallus ; when mature
they form a protuberance on the surface of the thallus and in section are seen to
be surrounded by a pericarp, as shown in Baardseth’s drawing (his fig. 41 Dp). The
tetrasporangia are also developed on the concave side of the upper thallus and can
be seen as dark patches. They do not protrude from the thallus. The spore mother
cells develop in accessory filaments (Fig. 76) from the lower cells of the cortex.
When mature (Fig. 77), the tetrads of spores measure up to 35 in diameter.

Baardseth commented at length on his reasons for including this species in the
genus Rhodoglossum and the generally unsatisfactory distinctions between the
genera Rhodoglossum, Gigartina and Inidaea.

CERAMIACEAE

CENTROCERAS CLAVULATUM (Agardh) Montagne in Explor. Sci. Algérie 1840-42,
Bot. I : 140 (1846).
Ceramium clavulatum Agardh in Kunth, Synops. Pl.: 2 (1822).

Slaughtered Seal Bay, below 15 cm. in rock pools, 22 Dec. 1955, G.I.S.S. 1507.
Midshipman Rock, mid-littoral, 18 Feb. 1956, G.J.S.S. 1518 ; same locality, occas-
ional in standing water in upper littoral, 18 Feb. 1956, G.J.S.S. 1520. Church Rock,
frequent on vertical rock face, 5 Mar. 1956, G.I.S.S. 1531, 1535. Isolda Rock,

upper-littoral pools, 6 Mar. 1956, G.J.S.S. 1553.
_ Widely distributed in warm and temperate waters.

This plant was found frequently on Gough Island intertwined with other algae.
MICROCLADIA ALTERNATA Baards. in Res. Norw. Sci. Exped. Tristan da Cunha 9g :

99, figs. 50 A, 5I C-E (1941).

Midshipman Rock, mid-littoral, 18 Feb. 1956, G.I.S.S. 1578.

Tristan da Cunha group of islands.

A small quantity of this species was found growing as an epiphyte on Corallina
officinalis at the Midshipman Rock. Female and tetrasporic plants are present
and agree well with Baardseth’s description and material.

RHODOMELACEAE

BOSTRYCHIA MIXTA Hook. & Harv. in Hook., Lond. Journ. Bot. 4 : 270 (1845).—
E. Post in Rev. Algol. 9 : 39 (1936).
Standoff Rock, on less-exposed part of the rock above the Corallina zone, 18 Feb.
1950, G.I.S.S. 1526.
Southern circumpolar temperate and cold waters, Japan, China.

B. mixta is a very common plant of southern shores where it frequently dominates
a community at or above high-tidemark, especially in places where there is a certain
amount of freshwater seepage. On Gough Island it was found only at Standoff
Rock, where it grew mixed with Rhizoclonium ambiguum, R. riparium forma validum

228 MARINE ALGAE OF GOUGH ISLAND

and Spongomorpha sp. Dr. Erika Post kindly confirmed the identification of this

plant.

HERPOSIPHONIA PANICULATA Baards. in Res. Norw. Sci. Exped. Tristan da Cunha
Q: 127, figs. 69 A, 71 A-C, 72 (1941). (Figs. 78, 79.)

Midshipman Rock, in mid-littoral zone on shiny olive-green deposit, 18 Feb. 1956,
G.I.S.S. 1515, 1517, 1518 ; same locality, in standing water in upper littoral,
18 Feb. 1956, G.I.S.S. 1520. Church Rock, vertical rock face, 5 Mar. 1956, G.I.S.S.
1535. Isolda Rock, in upper littoral on ridges between pools, 6 Mar. 1956, G.J.S.S.
1552.

Tristan da Cunha group of islands.

WOE
WS

SD WR
te YY
SR S CF
RQ WIE
WROTE
Q

Oe,

100u

19

78

Fics. 78,79. Herposiphonia paniculata Baards. : 78, ventral view of plant, y = rhizoid,
s = short branch, / = long branch ; 79, tetrasporic branchlet.

H. paniculata was a frequent epiphyte on Corallina officinalis. The plants
(Fig. 78) measure up to 2:5 cm. high. The main axis consists of a central cell
surrounded by 12-14 pericentral cells each about two to three times as long as wide.
Each segment of the main axis gives off a branch ; on every fourth segment are
alternating, ventral, long branches which repeat the structure of the main axis ;
on the intervening segments are simple branches up to 27 segments long and Ioop.
wide. This organization is characteristic of the genus. Tetrasporic (Fig. 79)
and carposporophytic plants are present ; the mature cystocarps on the latter grow
laterally on the simple branches and measure up to 350% wide X 500u long. The
tetraspores are borne in branchlets of the compound branches.

The Gough Island material agrees well with that described by Baardseth, who
points out that this plant may later prove to be within the variation range of one
of the other species of Herposiphonia with sexual organs occurring laterally on the
branchlets. In habit, the plants closely resemble the North American species

MARINE ALGAE OF GOUGH ISLAND 229

H. grandis Kylin and H. rigida N. L. Gardn., but they have consistently more
pericentral cells than either of these species.

LOPHURELLA sp. (Fig. 80.)

Church Rock, occasional amongst Cladophora radiosa on vertical rock face, 5 Mar.
1956, G.I.S.S. 1533.

It is unfortunately impossible to identify the species of this material with certainty
as the plants are incomplete, the ultimate branches being absent. However, as
the genus is predominantly sub-antarctic and antarctic, its presence on Gough
Island is of interest. The wiry plants (Fig. 80) measure up to 5 cm. high and

lcm

543 .
RA 80
Fic. 80. Lophurella sp. : habit of plant.

arise from a creeping, rhizomatous base. Tetraspores are present. The plants
belong to one of two species, L. hookerana (J. G. Agardh) Falkenb. or L. patula
(Hook. & Harv.) De Toni, but, without the terminal branchlets, it is impossible
to say which.

POLYSIPHONIA BOERGESENII Baards. in Res. Norw. Sci. Exped. Tristan da Cunha
g: 116, figs. 64 c, 65 B—D (1941).
West Point Reefs, pools below spray zone, 6 Mar. 1956, G.I.S.S. 1541.
Tristan da Cunha group of islands.

The Gough Island material of this species was found growing as an epiphyte on
Codium fragile in rock pools at West Point Reefs.

The material agrees exactly both with Baardseth’s description of the type and
with isotype material in the British Museum Herbarium except in the size of the

230 MARINE ALGAE OF GOUGH ISLAND

spermatangial branchlets. In the Gough Island material these are 140-180 long
whereas Baardseth describes them as being about 75y long. However, when I
measured the spermatangial branchlets in the isotype material, I found that these
were of the same order of size as those in the Gough Island plants ; it seems therefore
that Baardseth must have been measuring young branchlets.

POLYSIPHONIA HOWEI Hollenb. apud W. R. Taylor in Allan Hancock Pacif. Exped.
I2: 302, fig. 3 (1945).—Joly in Univ. Sao Paulo Fac. Fil. Ci. & Let. Bol. 217,
Bot. 14: 164, t. 13 figs. 5, 5a (1957).—W. R. Taylor, Mar. Alg. E. Trop. &
Subtrop. Coasts Amer. : 582 (1960). (Plate Ig A.)

Polysiphonia sp. Baards. in Res. Norw. Sci. Exped. Tristan da Cunha 9 : 119 (1941).

Midshipman Rock, mid-littoral, 18 Feb. 1956, G.I.S.S. 1517, 1518. Isolda Rock,
abundant on margins of mid-littoral pools, 6 Mar. 1956, G.J.S.S. 1549 ; same
locality, occasional in upper littoral on ridges between pools, 6 Mar. 1956, G.J.S.S.

1552.
Atlantic and Caribbean from North Carolina to Brazil, Tristan da Cunha, Pacific
from Panama to Colombia.

P. howei was collected from Midshipman Rock and from Isolda Rock, where it
grew in a low, brown turf on lower littoral rocks ; on Isolda Rock it actually domi-
nated a zone below Iridaea laminarioides and above Durvillaea antarctica. A
similar zone above the Durvillaea zone was reported on West Point Reefs and was
probably dominated by P. howe1, but no collection was made.

The plants from Isolda Rock (Plate 19 A) measure about 1 cm. high and have the
habit of a Lophostphonia with well-developed prostrate shoots, which have rhizoids
below and erect, sparingly branched filaments above. From the structure of the
apices of the prostrate shoots it can be clearly seen, however, that this plant belongs
to the genus Polysiphonia and not Lophostphonia ; the branches develop exo-
genously on all sides of the shoot and the dorsiventral habit is due to the fact that
the branches on the lower surface abort and rhizoids develop from the pericentral
cells on the lower side. The rhizoids are long, thin-walled, cut off from the peri-
central cell which produces them, and they end in a well-developed hapteron. The
erect shoots are strongly recurved at their apices and the apices bear dichotomously
branched trichoblasts. The filaments have 8-13, mostly 10-11, pericentral cells ;
the segments of the basal filaments measure about 130% wide xX 150 long, those
of the erect filaments 70-100n wide x 55-85yu long. The erect filaments branch
rarely when sterile but more abundantly when tetrasporic. Tetraspores are pro-
duced in the upper branches in somewhat spiral series ; there is one tetraspore
per segment. This material is in close agreement with Hollenberg’s and Taylor’s
descriptions of the species both in morphology and habitat. The plants from the
Midshipman Rock are very similar but have somewhat longer segments ; those of
the prostrate shoots are 100-1404 wide x 140-170y long. I do not think there is
any doubt however that they belong to the same species. The plants recorded by
Baardseth from the Tristan da Cunha group of islands as Polystphonia sp. are
also P. howet.

MARINE ALGAE OF GOUGH ISLAND 231

LOPHOSIPHONIA SCOPULORUM (Harv.) Womersley in Trans. R. Soc. S. Austral. 73 :
188 (1950).—Cribb in Univ. Queensl. Pap., Bot. 3: 138, t. I, t. 2 figs. 8-12
(1956). (Plate 19 B.)

Polysiphonia scopulorum Harv. in Trans. R. Irish Acad. 22 : 540 (1855).—Segi in Journ.
Fac. Fish. Pref. Univ. Mie 1 : 200, fig. 9, t. 3 fig. 7 (1951).
Polysiphonia villum J. G. Agardh, Sp. Gen. Ord. Alg. 2 (3) : 941 (1863).
Standoff Rock, on less-exposed part of the rock above the Corallina zone, 18 Feb.
1956, G.I.S.S. 1526.
Widespread in the Pacific Ocean, Australia, New Zealand.

This species was collected from Standoff Rock, where it formed part of a dense
turf together with Polysiphonia sp. The plants (Plate 19 B) are rather flaccid and
seldom measure more than 1-5 cm. high. They are composed of a prominent,
prostrate basal filament which gives off rhizoids mainly from its lower surface and
erect filaments mainly from its upper surface ; sometimes the positions are reversed.
The rhizoids are long and end in frilly haptera ; their contents are a continuation
of those of the parent pericentral cell. The filaments are ecorticate and have four
pericentral cells throughout. The basal segments measure 140-250 long x
c. 140u wide in the older parts and 70-gou long x c. 125y wide nearer the apex.
The erect filaments have segments 70-120u long xX I1oy wide and these in their
upper parts give off side branches with noticeably much shorter segments measuring
30-55u long xX 704 wide. Branching is quite frequent and in tetrasporic filaments
it is abundant. Tetraspores are borne in the side branches and there is one per
segment. Branching is endogenous ; no trichoblasts were seen.

The Gough Island plants agree well with isotype material in the British Museum
Herbarium and with Queensland material named by Dr. Cribb.

SPECIAL LITERATURE

BAARDSETH, E. (1941). The marine algae of Tristan da Cunha. Res. Norw. Sci. Exped.
Tristan da Cunha 9 : 1-174.

BORGESEN, F. (1939). Marine algae from the Iranian Gulf especially from the innermost part
near Bushire and the Island Kharg. Dan. Sci. Invest. Ivan 1 : 47-141.

CHAPMAN, V. J. (1956). The marine algae of New Zealand. Part I. Myxophyceae and
Chlorophyceae. Journ. Linn. Soc. Lond., Bot. 55 : 333-501.

DickiE, G. (1876). Contributions to the botany of H.M.S. ‘‘ Challenger”. XXIV. Marine
algae collected by Mr. Moseley at the Island of Kerguelen. Journ. Linn. Soc. Lond., Bot. 15:
43-47-

FREI, E., & PREston, R.D. (1961). Cell wall organization and wall growth in the filamentous
green algae Cladophora and Chaetomorpha. I. The basic structure and its formation.
Proc. R. Soc. Lond., Ser. B, 154 : 70-94.

Gain, L. (1912). La flore algologique des régions antarctiques et subantarctiques. 2me.
Expéd. Antarct. Frang., Bot. : 1-218.

HameEL, A., & Hamet, G. (1929). Sur l’hétérogamie d’une Cladophoracée, Lola (nov. gen.)
lubrica (Setch. et Gardn.). Compt. Rend. Acad. Sci. Par. 189 : 1094-1096.

HAMEL, G. (1931). Chlorophycées des cétes frangaises. (Fin.) Rev. Algol. 6: 9-73.

HAMEL, G., & LEMoOINE, P. (1952). Corallinacées de France et d’Afrique du Nord. Arch.
Mus. Nation. Hist. Nat. Par., Sér. 7,1 : 15-136.

Hariot, P. (1889). Algues. Miss. Sci. Cap Horn 5 : 3-109.

(1907). Algues. Expéd. Antarct. Frang., Bot., Alg. : 1-9.

232 MARINE ALGAE OF GOUGH ISLAND

Hooker, J. D. (1845-47). Botany of Fuegia, The Falklands, Kerguelen’s Land, etc. The
Botany of the Antarctic Voyage. 1. Flora Antarctica : 209-574.
Isaac, W. E. (1957). The distribution, ecology and taxonomy of Porphyra on South African
coasts. Proc. Linn. Soc. Lond. 168 : 61-65.
Jory, A. B. (1957). Contribuig¢ao ao conhecimento da flora ficol6gica marinha da Baia de
Santos e Arredores. Univ. Sdo Paulo Fac. Fil. Ci. & Let. Bol. 217, Bot. 14 : 1-199.
Knox, G. A. (1960). Littoral ecology and biogeography of the southern oceans. Proc. R.
Soc. Lond., Ser. B, 152 : 577-624.
LeEvrinG, T. (1938). Verzeichnis einiger Chlorophyceen und Phaeophyceen von Siidafrika.
Act. Univ. Lund., Ny Foljd, Avd. 2, 34 (9) : I-25.
(1953). The marine algae of Australia. I. Rhodophyta: Goniotrichales, Bangiales
and Nemalionales. Ark. for Bot., Ser. 2, 2 : 457-530.
(1956). Contributions to the marine algae of New Zealand. I. Rhodophyta : Gonio-
trichales, Bangiales, Nemalionales and Bonnemaisoniales. Ark. for Bot., Ser.2,3: 407-432.
(1960). Contributions to the marine algal flora of Chile. Act. Univ. Lund., Ny Foljd, Avd.
2, 56 (10) : 1-85.
Manza, A. V. (1937). The genera of articulated corallines. Proc. Nation. Acad. Sci. Wash.
23 : 44-48.
(1940). A revision of the genera of articulated corallines. Philipp. Journ. Sci. 71 :
239-310.
Papenrfuss, G. F. (1960). On the genera of the Ulvales and the status of the order. Journ.
Linn. Soc. Lond., Bot. 56 : 303-318.
Post, E. (1959). Weitere Daten zur Verbreitung des Bostrychietum VII. Ayvch. Protistenk.
103 : 489-500. :
StnHA, J. P. (1958). Chromosome numbers and life-cycles in members of Cladophorales.
Brit. Phyc. Bull. 1 (6) : 24-27.
SKOTTSBERG,C. (1923). Botanische Ergebnisse der Schwedischen Expedition nach Patagonien
und dem Feuerlande 1907-09. IX. Marine algae 2. Rhodophyceae. K. Svensk. Vetensk-
apsakad. Handl., Ny Foljd, 63 (8).
(1941). Communities of marine algae in subantarctic and antarctic waters. K. Svensk.
Vetenskapsakad. Handl., Ser. 3, 19 (4).

STEPHENSON, T. A. & A. (1954). Life between tide-marks in North America. IIIa. Nova
Scotia and Prince Edward Island : description of the region. Journ. Ecol. 42 : 14-70.
SUNESON, S. (1943). The structure, life-history and taxonomy of the Swedish Corallinaceae.

Act. Univ. Lund., Ny Foljd, Avd. 2, 39 (9) : 1-66.
WomersLey, H. B.S. (1956). A critical survey of the marine algae of southern Australia. I.
Chlorophyta. Austral. Journ. Mar. Freshw. Res. 7 : 343-383.

Bull. B.M. (N.H.) Bot. 3, 5 PLATE 16

B Cc

A. Theshore at Isolda Rock. B. Colpomenia sinuosa (Roth) Derb. & Solier, habit of plant
(x2). C. Corallina goughensis Y. M. Chamberlain, habit of plant ( 2).

Bull, B.M. (N.H.) Bot. 3, 5 PLATE 17

Dermatolithon nodulosum Y. M. Chamberlain. A. Habit of plant (x). B. Detail of
nodules (x11). C. Transverse section through nodule with tetrasporic conceptacles.

Bull. B.M. (N.H.) Bot. 3, 5 PLATE 18

Lithophyllum sp.

Lithothamnion neglectum (Foslie) Foslie. A. Habit of G.I.S.S. 1548 (x $), B. Habit of
G.I.S.S,. 1553 (x *). CC. Vertical section through G.I.S.S. 1507.

oe
i]
Te | "= _ = owe : = i. - x a «tt a a ue a, a = or
i) ;
= : a a : 7 ae a) Xi : _ :
> _ 7 _ - 7 7 : i ea | ;
7 - 7 : - : : 7 a
| | : eas

Bull. B.M. (N.H.) Bot. 3, 5 PLATE 19

|

‘A ) ~

B

A. Polysiphonia howei Hollenb. Habit of plant (x10). B. Lophosiphonia scopulorum
(Harv.) Womersley. Habit of plant (x 12).

tf

q
THE CEYLON SPECIES
OF ASPLENIUM

W. A. SLEDGE

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 3 No. 6
LONDON: 1965

: af
:

Wise ere sre
Apia

eae at
reeds

Tee CEYLON SPECIES OF ASPLENIUM

N.S OCT 1965
— x

* é
AL Wes

BY
W.. A. SLEDGE

(University of Leeds)

Xu/

J

Ph. 233-277 ; 3 Text-figures ; Plate 20

BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
BOTANY Vol. 3 No. 6
LONDON : 1965

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THE CEYLON SPECIES OF ASPLENIUM
By W. A. SLEDGE

ASPLENIUM is the largest genus of ferns in Ceylon. Many of the species are well
defined and easy to recognize ; others are highly polymorphic. No preliminary
discussion of the former group is required. The taxonomic, nomenclatural and
distributional points which arise in connexion with some of them are discussed
under the appropriate species. As regards the more polymorphic species my aim
has been to define the limits of their variability without assigning names to the
diverse forms which they comprehend. As this procedure has led in some cases to
the referring of rather heterogeneous assemblages of seemingly different plants to
single species, some discussion of the problems involved and some justification of
the method adopted are required.

In recent years a rapidly growing body of research work has demonstrated the
existence of cytological complexities in numerous genera of ferns. Asplenium is
notable for the wide range of polyploidy shown within the genus and many hybrids
have been described. Chromosome counts have been made (Manton & Sledge in
Phil. Trans. R. Soc. Lond., Ser. B, 238 : 127-185 (1954)) of about half the Ceylon
species of Aspleniwm and these have established the presence there of diploid,
triploid, tetraploid, octoploid and dodecaploid plants. More than one grade of
polyploidy exists within some species and clear cytological evidence of hybridity was
obtained in one of the stock plants examined. Having regard to the necessarily
limited number of plants investigated in a project designed to explore the cytology
of the fern flora of Ceylon in general and not aimed at any one genus in particular,
it is more than likely that hybrids are frequent and that apogamy and other grades
of polyploidy exist both within unworked species and within species at present
known only from counts based on single plants. While most of the species appear
to be tetraploid, A. affine is represented by octoploid and dodecaploid races. Three
plants of A. aethiopicum were also found to be octoploid in Ceylon. In Africa
tetraploid, octoploid and dodecaploid races of A. aethtopicum are known to occur
and the morphology of tetraploid plants is closely similar to that of some Ceylon
plants.

Illustrations are given in our paper (Manton & Sledge, loc. cit., fig. 17) of fronds
from octoploid and dodecaploid plants of A. affine. The differences displayed
would be sufficient to warrant taxonomic distinction if all plants agreed either with
the one or the other ; but in fact a series of intermediate forms exists such that no
useful separation or subdivision can be drawn. A dodecaploid race is most likely
to have involved the participation of some other species in its origin, and it would be
expected that some character or characters could be found whereby the two races
might be distinguished. Such hereditable differences might well be found if large

236 THE CEYLON SPECIES OF ASPLENIUM

suites of living plants were assembled and kept under constant observation, but the
taxonomic difficulty in such species as A. affine and A. aethiopicum lies in the fact
that both show great inherent plasticity and the range of phenotypic modification
produced by this plasticity is closely paralleled and overlapped by the range of
variation produced by genotypic diversity.

Evidence of this inherent plasticity is shown in the accompanying silhouettes
(Figs. 1, 2) which show fertile fronds taken from wild specimens of A. aethiopicum
and A. affine and fronds taken from the same rhizomes after cultivation at Kew for

Fic. 1. Asplenium aethiopicum (Burm. f.) Becherer. The left-hand frond from a wild
plant (P.222) collected in Ceylon in 1951 ; the right-hand frond taken from the same
rhizome after one year in cultivation at Kew. Both x }.

one year and five years respectively. Apart from the size, the variable characters
include degree of pinnation and shape of pinnules in both species, the presence or
absence of proliferating buds on the rhachis in A. affine (absent in the frond taken
from the wild plant but present on that produced in cultivation, both on the main
rhachis and on the pinna rhachides, though too small to show in the silhouette),
and the degree of development of scales on the rhachis and pinnae of A. aethiopicum
(densely woolly-scaly in the frond taken from the wild plants but very sparsely scaly
in the frond produced in cultivation). The differences in gross morphology between
the two fronds of each pair considerably exceed those which have served, sometimes
rightly and sometimes wrongly, to distinguish between species.

THE CEYLON SPECIES OF ASPLENIUM 237

The crucial point in the taxonomic treatment of such polymorphic macrospecies
is that discontinuity of variation must determine the limits of species. The lower
limit to the degree of morphological difference that can be accepted as providing a
specific distinction is that by which a competent taxonomist can with certainty
discriminate between two groups. But there is no upper limit to the morphological
variability that can be admitted within a species if not accompanied by at least some
degree of reproductive isolation. That some species are larger and more poly-
morphic than others is an inescapable fact.

Fic. 2. Asplenium affine Swartz. The left-hand frond from a wild plant (P.52) collected

in Ceylon in 1951 ; the right-hand frond taken from the same rhizome after five

years in cultivation at Kew. Both x 3.

The evidence supplied by these two plants is alone sufficient to show that the
great differences displayed are of no value for taxonomic purposes. Where so wide
a response to age and environment is shown by individual plants, it is clearly im-
possible without full-scale cytotaxonomic investigations to define the taxa present
within each macrospecies ; and, indeed, even then it is by no means impossible
that the phenotypic expression of genetic differences may be too much at the mercy
of environmental influence for it to be of any taxonomic value, since it may well be
impossible to assess differences with confidence in a dried specimen or even in the
majority of living specimens. Most taxonomists feel dissatisfied at leaving unnamed
a series of grossly divergent morphological types ; but where the pattern of variation

238 THE CEYLON SPECIES OF ASPLENIUM

is so continuous, with many variables and little or no correlation, the division of the
species into a series of subspecific units, designed to cover the range of variation in
such a way that they will be recognizable to some one other than the author who
establishes them, is practicable only for a limited number of specimens since the
boundaries between such units must be purely arbitrary. To assign Latin
names to such arbitrary divisions is not only worthless : it is potentially harmful in
that later taxonomists, impressed by the great apparent difference of a specimen to
which some infraspecific rank has been given, may raise it to independent specific
status and in so doing create a binomial which divorces it from the species to which
it properly belongs.

I have discussed elsewhere (Kew Bull. 15 : 401 (1962)) the taxonomy of A. affine
and shown that the supposedly distinct species to which the name A. sfathulinum
has been widely applied is a clear case of two names covering a single species, since
the distinctions were based on variable, non-correlated characters. The most
extreme variant, which I distinguished as a forma, had been treated by Hooker
(Sp. Fil. 3 : 169 (1860)) as typical A. affine. Owing to acceptance of a tradition
concerning the different geographical ranges of the two supposed species, the applica-
tion of the names A. affine and A. spathulinum as accorded to most specimens in
herbaria is determined usually by the provenance of the specimen concerned rather
than by its morphological characters. The fact that the species A. affine is now
known from cytological evidence to be an unresolved species complex affords no
justification whatever for the perpetuation of a name originally given to a form
embraced within its own known amplitude of variation. In the case of A. aethiopi-
cum I have named no variants, though earlier botanists sometimes gave specific rank
to one of its Ceylon forms, to which they incorrectly applied the name A. laserpitii-
folium. A detailed cytogenetic investigation of the African forms of A. aethiopicum
is at present being pursued by Mr. A. F. Braithwaite and his findings may be expected
to throw some light on the taxonomy of this species as represented in Ceylon.

I have referred only to A. affine and A. aethiopicum in this discussion since for
them direct evidence of plasticity can be supplied. But the same considerations
apply to other polymorphic species, of which A. evectum and A. falcatum are examples.
Only a brief reference to these species need be made here since they are fully dis-
cussed later on. In the case of A. evectum, species, varieties and forms have been
founded on variations in pinna form which I consider of no taxonomic signifi-
cance. Such taxa have frequently been described by authors with no knowledge of
the living plants or the range of variation displayed in wild populations, their
differences being more a reflection of the limited numbers of herbarium specimens
which were available for study than an expression of valid taxonomic distinctions.
Selected examples might be taken from any large population of A. erectuwm in Ceylon
that would display considerable range in frond and pinna shape and would cover
forms to which some authors have assigned independent rank. Such differences
I consider examples of intraspecific variation of a fluctuating nature. In the case of
A. falcatum, plants with bipinnate fronds are of frequent occurrence in Ceylon and
these have been confused in the past with other species. No evidence is available

THE CEYLON SPECIES OF ASPLENIUM 239

to show if such plants are able, like Diplazium dilatatum, to produce both simply
pinnate or bipinnate fronds according to age or environmental conditions, or whether
they are fixed varieties. They are, however, so widely different in appearance
from the usual simply pinnate condition that nomenclatural recognition is here very
desirable as a means of referring such plants to their correct species.

Thwaites (Enum. Pl. Zeyl.: 383 (1864)) recognized twenty species of Asplenium.
As with all Ceylon ferns, his arrangement closely followed Hooker’s account in the
Species Filicum, both in the matter of classification and nomenclature. Beddome’s
account of the genus in his Handbook to the Ferns of British India, Ceylon and the
Malay Peninsula (1883) hardly differs, such changes as are made largely following
those introduced in the Synopsis Filicum. Little detailed attention has since been
paid to Ceylon ferns. The twenty-two species enumerated in Willis’s Catalogue of
the Plants of Ceylon (1911) represent a revised list compiled from Christensen’s
Index Filicum.

In the following account I recognize twenty-five species. A. macrophyllum, which
was listed by Beddome and Willis, I am unable to maintain as a distinct species in
Ceylon whatever its status elsewhere. One of the four additional species,
A. inaequilaterale, was recognized by Beddome under the name A. lunulatum var.
trapeziforme. A. obscurum and A. pellucidum have not previously been recorded
from Ceylon and one new species, A. disyunctum, is described.

Most of the Ceylon species of Asflenium are montane plants, commonly growing
on rocks and trees in jungle and forest, or as ground species. The only species
common in the lowlands are A. nidus and A. falcatum. Two species, A. pellucidum
and A. nitidum, are known from Ceylon only from single specimens in herbaria.

The most widely distributed of the Ceylon species of Asplenium is A. aethtopicum
which, in one or other of its many forms, is found in tropical America, Africa, Asia,
Australia and Polynesia. A. formosum and A. inaequilaterale also grow in tropical
America and Africa but in Asia are confined to peninsular India and Ceylon. Species
which range eastwards only from Ceylon and India are A. ensiforme, A. chetlosorum,
A. indicum, A. nitidum, A. decorum and A. tenuifolium. Of these the second, third,
fifth and sixth reach the Philippine Islands. The following widely distributed
species range both westwards to Africa or the Mascarene Islands and eastwards to
the Philippines or Polynesia: A. nidus, A. normale, A. unilaterale, A. obscurum
(Madagascar to China), A. erectum, A. tenerum, A. pellucidum, A. falcatum, A. affine
and A. varians.

Of the species with more restricted ranges, A. zenkeranum, A. decrescens and
A. serricula are confined to Ceylon and South India, the last-named species also
occurring in the Philippine Islands. A. gardneri is known only from Ceylon and
Sumatra. A. longipes and A. disjunctum are endemic to Ceylon. Apart from the
two endemics, A. gardnert and A. decorum are the only other Ceylon species of
Asplenium which are not also found in India.

My grateful thanks are due to the Directors and Curators of the following herbaria

240 THE CEYLON SPECIES OF ASPLENIUM

for access to, or for the loan of, specimens in their charge. Abbreviations used in
the citation of specimens are those adopted in the Index Herbariorum.

BM = British Museum (Natural History), London.

CGE = Botany School, University of Cambridge.

E = Royal Botanic Garden, Edinburgh.

G = Conservatoire et Jardin Botaniques, Geneva.

GH = Gray Herbarium of Harvard University, Cambridge, Mass.

GL = Department of Botany, University of Glasgow.

K = Royal Botanic Gardens, Kew.

PDA = Department of Agriculture, Peradeniya.

SING = Botanic Gardens, Singapore.

US = United States National Museum (Department of Botany),
Smithsonian Institution, Washington, DC.

W = Naturhistorisches Museum, Vienna.

In the citation of specimens the numbered ones distributed by Thwaites have
been listed first since sets of these specimens are to be found in many herbaria and
the numbers have been widely quoted. Statements as to localities where these
plants were collected rarely accompany the specimens and the data which have been
pencilled on many of the sheets in the Peradeniya collection are indicative of localities
whence the species had been recorded rather than of localities where the actual
specimens on the sheets were collected. Since errors of identification occur, the
localities are not therefore always reliable. Earlier sets of specimens were distributed
by Gardner and of these the Cambridge sheets bear his own annotations giving name,
place of origin and date of collection. Other gatherings by Gardner bear numbers
only.

ASPLENIUM L.

KEY TO THE SPECIES
Fronds simple :
Veins connected by a marginal strand ; fronds large. ; 1. A. nidus
Veins free ; fronds small to medium-sized. : : . 2. A. enstforme

Fronds pinnate, bipinnate or tripinnate :
Fronds simply pinnate :

Pinnae 1-2 cm. long ; stipes black and polished ; rhizome short, fronds tufted :
Upper margin of pinnae deeply cut : ; : . 3.A. formosum
Upper margin of pinnae crenate or dentate. : ; 4. A. normale

Pinnae more than 2 cm. long :

Stipe black and polished ; rhizome creeping :
Upper margin of pinnae deeply cut, sori confined to the lobes
5. A. cheilosorum
Upper margin of pinnae serrate, sori not on the lobes 6. A. unilaterale
Stipe green or, if dark, not glossy and polished :
Pinnae dimidiate, the lower side cut away to the costa ; rhizome long-
creeping . 3 ; ; i ; : . 7. A. obscurum

THE CEYLON SPECIES OF ASPLENIUM 241

Pinnae subequal-sided or the lower side cuneate, or if dimidiate then
rhizome erect :
Rhizome short, fronds tufted :
Pinnae up to 5 cm. long :
Fronds decrescent . ‘ : ; . ‘ 8. A. evectum

Fronds not decrescent :
Veins forked :

Stipe and rhachis scaly. : ; . 18. A. indicum
Stipe and rhachis not scaly : : g. A. tnaequilaterale
Veins simple (except for lowermost acroscopic ones) :
Pinnae toothed : : é ; : 10. A. tenerum
Pinnae cut to the costa into linear lobes . 25. A. decorum
Pinnae more than 5 cm. long :
Fronds decrescent . : ; 2 : 11. A. pellucidum
Fronds not decrescent :
Fronds gemmiparous . , 5 ; 12. A. zenkeranum

Fronds not gemmiparous :
Stipe green :
Pinnae 0:5—2 cm. broad, widest at the middle 13. A. serricula
Pinnae 2-4 cm. broad, widest near the base 14. A. longipes

Stipe black ; , 17a. A. falcatum var. falcatum
Rhizome creeping, fronds more or less distant :
Fronds gemmiparous ; sori diverging . . 15. A. gardnert

Fronds not gemmiparous ; sori parallel with the costa
16. A. decrescens
Fronds bipinnate or tripinnate :
Pinnules ovate, spathulate or narrowly rhomboid, lobed or dentate but not

divided into narrow segments: -
Medium to large ferns ; texture of frond firm :
Stipe and rhachis quite glabrous : : : . 19. A. nitidum

Stipe and rhachis scaly :
Veins very close ; fronds not gemmiparous :
Stipe and rhachis (at least when young) woolly with hair-pointed
scales ; rhizome creeping. : ; 20. A. aethiopicum
Stipe and rhachis not woolly-scaly ; rhizome short, fronds tufted
17b. A. falcatum var. bipinnatum
Veins spaced ; fronds often gemmiparous ; : 21. A. affine
Small fern ; texture of frond thin, herbaceous ; : 22. A. varians
Pinnules linear, I-2 mm. broad, or divided into narrow segments :
Fronds gemmiparous :
Fronds bipinnate-tripinnatifid . , : . 23. A. disjunctum
Fronds tripinnate-quadripinnatifid . : : 24. A. tenuifolium
Fronds not gemmiparous ; pinnules linear. : . 25. A. decorum

242 ERE CEYLON SPECIES (OF ASPLEENTUM

1. Asplenium nidus L., Sp. Pl. 2: 1079 (1753).

Thamnopteris nidus (L.\) C. Presl in Abhandl. K. Béhm. Ges. Wiss., Folge 5, 6 : 428 (1851).
—Bedd., Handb. Ferns Brit. Ind. : 137 (1883).

Rhizome stout, erect, apex clothed with lanceolate, attenuate, brown scales
bordered by filamentous appendages. Fronds tufted, in a rosette, simple, variable
in length and breadth, commonly 100 x ro cm. but often larger or sometimes smaller,
base narrowed into a short stipe, apex acute or acuminate, margins entire, costa
prominently raised beneath ; veins simple or once, sometimes twice, forked, uniting
to form a continuous intramarginal vein ; texture stiff, subcoriaceous. Sori narrow,
extending from the costa 4-4 the way to the margin. Spores plano-convex, 33-39 X
24-27, with an undulate perispore wing and anastomosing surface folds.

CEYLON : Thwaites C.P. 3267 (E; PDA). On trees, hotter parts of island
(Rattotta, alt. 300 m., in Herb. Edin.), Naylor Beckett 299 (BM ; E ; GH). Nagolla,
Matale, Apr. 1858, May 1858, Herb. Brodie (E). Hulangamuwa, Matale,
May 1858, Herb. Brodie (E). Kelani Valley, Freeman 324 A, 325 B (BM).
Summit of Ritigala (alt. 750 m.), 22 Mar. 1905, Willis (PDA). Terrestrial in forest
on slopes of Ritigala, 600 m., 14 Jan. 1951, Sledge 871 (BM). Opanake between
Balangoda and Ratnapura, epiphytic in forest, 150 m., 3 Jan. 1951, Sledge 800 (BM).
Anderson (E). Mrs. Chevalier (BM). Ferguson (GH ; PDA ; US 815500). Koenig
(BM). Macrae (E). Wall (E; GL).

Common in the low country up to 300 m. and ascending to 750 m.; usually
epiphytic but sometimes terrestrial.

Tropics of the Old World.

2. Asplenium ensiforme Wall. ex Hook. & Grev., Ic. Fil. : t. 71 (1829).

Asplenium ensiforme Wall., Numer. List : 8, n. 200 (1829), nom. nud.

Rhizome short, suberect, bearing narrow, attenuate, dark-brown scales at its
apex and at the base of the stipes. Fronds tufted, 15-35 X I-3cm., entire, narrowly
oblong or elliptic, narrowed at the base into a stipe 1-2 cm. long, gradually acuminate
above, margins entire or slightly repand, both surfaces glabrous save for a few
filiform scales on the lower part of the midrib beneath ; veims immersed, mostly
once forked, free; texture stiff, subcoriaceous. Sori spreading along the veins from
near the costa nearly to the margin. Spores reniform, 42-48 x 24-30, with a
wide undulate perispore wing and anastomosing surface folds.

CEYLON: Thwaites C.P. 1334 (BM; GH; K ; PDA—data on PDA sheet :
Ramboda, Sept. 1847, Oct. 1853, Gardner). Hantane range, on trees in forests,
Aug. 1844, Gardner 1155 bis (CGE). Central Province, go00-1,200 m., Naylor
Beckett 268 (BM ; GH). Knuckles Mt. near Kandy, epiphytic in forest, 1,650 m.,
30 Jan. 1954, Sledge ro81 (BM ; K). Le Vallon Estate, on rocks and trees in jungle,
1,500 m., 9 Feb. 1954, Sledge 1112 (BM ; K ; US). Ramboda-Maturata track, on
rocks and trees by stream in forest, 1,920 m., 17 Mar. 1954, Sledge 1298 (BM; K).
1899, Anderson (E). Ferguson (PDA ; US 815480). Walker (K). Wall (E ; GL; K).

On rocks and trees in forests above 1,000 m.

THE CEYLON SPECIES OF ASPLENIUM 243

Ceylon, South India, Himalaya from Punjab to Assam, south China, Burma,
Thailand, Indochina.

3. Asplenium formosum Willd. in L., Sp. Pl. ed. 4, 5 : 329 (1810).— Hook.,
Sp. Fil. 3 : 143 (1860).—Bedd., Handb. Ferns Brit. Ind. : 152 (1883).

Rhizome short, erect, scaly at the apex, scales linear, entire, black with brown
margins. Stipes 1-5 cm. long, dark-purpfle to black, naked, polished ; rhachis of the
same colour with a very narrow wing on each side which is continuous to the base of the
stipe. Lamina simply pinnate, up to 30 cm. long, 2 cm. wide, with 30-40 or more
pairs of pinnae, the lower ones reduced in size ; pinnae subsessile, horizontal,
dimidiate, upper base truncate to broadly cuneate, about half of the lower side cut
away, upper margin and distal part of lower margin deeply incised, the proximal
lobes mostly bifid ; veins immersed, forked or simple ; texture firm-herbaceous.
Sori short, oblique, I-3 per pinna and usually confined to the lower side of the costa.
Spores light-brown, 33-36 Xx 24u, plano-convex with a perispore wing and surface
folds.

CEYLON : Thwaites C.P. 3487 (BM ; E ; GH; K ; PDA—data on PDA sheets :
Galagama, Apr. 1856; East Matale, on rocks in river bed). Lagalla, 360 m.,
Naylor Beckett 26 (E; GH). Between Corbet’s Gap and Kaikawala, 750 m., Aug. 1956,
Abeywickrama 346 (Herb. Univ. Ceylon). Weragamtota, Central Province, on
rocks by stream in jungle, 450 m., 10 Jan. 1954, Sledge 957 (BM). Ferguson (GH ;
PDA). Henderson (US 418661-2). Wall (GL).

On rocks by streams in forests up to 1,000 m.; confined to the eastern and northern
edge of the central massif.

Tropical America from Mexico to Peru, West Indies, tropical Africa, Portuguese
East Africa, Madagascar, Ceylon, South India (Nilgiri, Palni and Anaimalai Hills).

This is one of a group of ferns confined, in Ceylon, to the transitional region be-
tween the wet and dry zones fringing the eastern and northern edges of the central
mountain massif. Athyrium hohenackeranum and Anisocampium cumingianum are
other species which show the same distribution.

Asplenium formosum has a similar distribution to A. inaequilaterale, ranging from
tropical America eastwards across Africa but reaching Asia only in peninsular India
and Ceylon.

4. Asplenium normale D. Don, Prodr. Fl. Nepal. : 7 (1825).

Asplenium minus Blume, Enum. Pl. Jav. 2: 183 (1828).

Asplenium multijugum Wall., Numer. List : 8, n. 207 (1829), nom. nud.

Asplenium opacum Kunze in Linnaea 24 : 261 (1851).

Asplenium pavonicum Brackenr. in U.S. Expl. Exped. Wilkes : 150, t. 20 fig. 1 (1854).

Asplenium multijugum Wall. ex Mett. in Abhandl. Senckenb. Naturforsch. Ges. 3 : 179
(1859).—Hook., Sp. Fil. 3 : 139, t. 188 B (1860).

Rhizome erect, the apex clothed with narrow brown scales. Stipes numerous,
tufted, 3-15 cm. long, wiry, polished, dark-chestnut to purple-black ; rhachis glossy
like the stipe, grooved above. Lamina pinnate, up to 30 cm. long, 2-4 cm. broad,

244 THE CEYLON SPECIES OF ASPLENIUM

linear-oblong with numerous (15-45) pairs of close-placed, sessile, patent pinnae,
and sometimes proliferous from a bud formed distally on the rhachis, upper pinnae
reduced in size, the lowermost scarcely reduced, often deflexed ; pinnae 0-75-2 x
0:5-0'75 cm., dimidiate, obtuse, upper base truncate, more or less parallel to the rhachis,
slightly auricled with the auricle sharply angled at the tip, lower base very obliquely
cuneate forming a more or less horizontal line 4-4 the length of the pinna, upper
margin and apex crenate-dentate, both surfaces glabrous ; one or more veins above
the costa forked, those below the costa simple ; texture stiff-herbaceous. Sori
short, variable in number, often one on the auricle and 2-3 others obliquely placed
above the costa and 1-2 below and more or less parallel with the lower margin.
Spores plano-convex, 30-36 x 24-27u, with a rather broad perispore wing and
anastomosing surface folds.

CEYLON: Thwaites C.P. to05 (BM; E; GH; K; PDA). Gardner 1073
(CGE ; E ; K—the CGE sheet labelled Nuwara Eliya, in dry forests, Sept. 1844).
Heights above Peradeniya, Gardner (K). Kandy, rocks, Macrae 287 (K). Pussalawa,
1855-57, 1871, Sall and Randall in Rawson 3220/369/298 (BM). Central Province,
750-1,200 m., Naylor Beckett 213 (E; GH). Adam’s Peak, 1,350 m., 14 Feb. 1908,
Matthew (K). The Hope, Hewahetta, 25 May 1862, Herb. Brodie (E).
Nuwara Eliya, Mrs. Chevalier (BM). Same locality, Freeman 134 A, 135 B, 136 C,
137 D (BM). Same locality, 1,950 m., 24-27 Feb. 1954, Schmid 1354, 1360 (BM).
Horton Plains, 2,270 m., 7-8 Mar. 1954, Schmid 1395 (BM). Passara, patana,
660 m., 1897, Pearson 407 (CGE). Corbet’s Gap, roadside through secondary
jungle, 1,200 m., 9 Dec. 1950, Sledge 562 (BM). | Above Ramboda, Robinson (K).
Ramboda Pass, on mossy rock by stream in jungle, 1,650 m., 17 Dec. 1950, Sledge 647
(BM). Ascent of Mt. Pedrotalagalla, in shady ground by side of track, 2,025 m.,
26 Dec. 1950, Sledge 726 (BM). Near Hakgala, on ground in shady places, 1,650 m.,
23 Dec. 1950, Holttum 39164 (SING). Hakgala, by stream in jungle, 1,650 m.,
27 Dec. 1950, Sledge 738 (BM). Mt. Namunukula, terrestrial in forest, 1,800 m.,
24 Feb. 1954, Sledge 1194 (BM). Ramboda-Maturata track, terrestrial in forest,
1,920 m., 17 Mar. 1954, Sledge 1297 (BM). Parawella Falls, Kandapola near Nuwara
Eliya, 1,425 m., 19 Mar. 1954, Sledge 1335 (BM). 1899, Anderson (E). Ferguson
(GH ; PDA; US 815505). Gardner 25 (BM). Macrae (E). 12 Mar. 1819,
Moon 482 (BM). Palliser (GH). Wall (E ; GH; GL).

Common in forests above 1,200 m.

Tropical East Africa and Madagascar, Mauritius, Ceylon, South India, Himalaya
from Nepal to Assam, south China, Formosa, Indochina, Malaya, Sumatra, Java,
Borneo, Philippines, New Guinea, Hawaii.

A. normale is like A. erectum in appearance but is distinguished by the dark,
glossy, polished stipe and rhachis. In A. erectum the stipe and rhachis may be
grey or dark-coloured but are never glossy and polished, and a slender green wing
borders each side of the groove on the upper surface of the rhachis. The margins
of the pinnae are more shallowly incised in A. normale, and the auricles (often
weakly developed) are sharply angled at the tip.

THE CEYLON SPECIES OF ASPLENIUM 245

I have examined the type specimen of Blume’s A. minus from Java. It is the
same as A. normale and has been so identified by Rosenstock.

Kunze admitted to not having seen examples of Don’s species when he described
A. opacum. There is an authentic example of the latter at Kew. It differs only
in its somewhat larger and broader pinnae, and is matched by some Ceylon forms.

5. Asplenium cheilosorum Kunze ex Mett. in Abhandl. Senckenb. Naturforsch.
Ges. 3 : 177, t. 5 figs. 12, 13 (1859).
Asplenium heterocarpum Wall., Numer. List : 9, n. 218 (1829), nom. nud.
Asplenium heterocarpum Wall. ex Hook., Sp. Fil. 3 : 132, t. 175 (1860).—Bedd., Handb.
Ferns Brit. Ind. : 153, t. 77 (1883).

Rhizome creeping, clothed with narrow, dark, tapering, acute scales. Stipes up
to 20 cm. long, dark-purple, polished, scaly at the base, otherwise glabrous ; rhachis
of the same colour, naked, glossy. Lamina up to 35 cm. long, 4—5 cm. broad, linear
in outline with numerous (up to 40 or more) pairs of pinnae, the lower ones very
shortly stalked ; pinnae 2-3 X 0-5-0°8 cm., dimidiate with nearly the whole of the
lower margin cut away, upper base broadly cuneate to truncate, upper margin and
extremity of lower margin lobed, the lobes mostly bifid ; veins once forked, the two
branches passing one to each tooth of a lobe ; texture thin, membranous. Sori
short, confined to the lobes and mostly to the distal fork only of each lobe ; indusium
thin, brown. Spores reniform to plano-convex, 42-51 Xx 30—-33y, with a rather
broad perispore wing and many reticulately anastomosing surface folds.

CEYLON : Thwaites C.P. 1006 (GH ; PDA—data cited on the PDA sheet :
Nuwara Eliya, Jan. 1847, Gardner ; Hewahetta, Oct. 1852 ; Pelagalla, Oct. 1853).
Thwaites C.P. 1363 (BM). Central Province, damp forests, Naylor Beckett 28 (BM).
Ramboda Pass, Freeman 149 B (BM). Nuwara Eliya, Freeman 148 A (BM). Near
Hakgala, on rocks by stream, 1,650 m., 27 Dec. 1950, Holttum 39202 (SING).
Dooroomadella, Matale, Dec. 1860, Herb. Brodie (E). Oodawella near
Kandy, in jungle above tea plantation, 1,050 m., 8 Dec. 1950, Sledge 521 (BM).
Corbet’s Gap, in secondary jungle, 1,320 m., 12 Dec. 1950, Sledge 556 (BM).
Hoolankande, terrestrial in shady forest, 1,350 m., 20 Jan. 1954, Sledge ror2 (BM).
1899, Anderson (E). 1828, Emerson ex Herb. Greville (E). Ferguson (GH ;
US 815493, 815494). Gardner 1074 (K). Gardner 1076 (BM). Herb. Harland 183
in Herb. Hance (BM). Robinson 57 (K). Wall (E ; GH ; US 418666).

In wet forests above 900 m.

Ceylon, North and South India, south China, Burma, Indochina, Malaya, Borneo,
Philippines, Formosa, Japan.

6. Asplenium unilaterale Lam. in Encycl. Méth., Bot. 2: 305 (1786).
Asplenium resectum Smith, Pl. Ic. Ined. 3 : t. 72 (1791).—Hook., Sp. Fil. 3: 130 (1860)
excl. parte.
Asplenium erythrocauion Blume, Enum. Pl. Jav. 2 : 183 (1828).
Asplenium laetum Wall., Numer. List : 8, n. 209 (1829), nom. nud.
Asplenium emarginatodentatum Zenker apud Kunze in Linnaea 24 : 263 (1851).

Rhizome creeping, slender, scaly at the apex, scales narrow, brown, margins entire.
Stipes 10-20 cm. long, dark-castaneous to black, glossy, naked ; rhachis of the same

240 THE CEYLON SPECIES OF ASPLENIUM

colour, shining, grooved above with a narrow green wing on each side of the groove.
Lamina simply pinnate, very variable in size, 15-35 5-12 cm., narrowly or broadly
oblong or oblong-lanceolate, the lowermost pinnae not or only slightly reduced in
size, often somewhat reflexed ; pinnae in 15-30 pairs, very shortly stalked,
patent, dimidiate, commonly 2-3 x 0:5 cm., sometimes up to 7 X I°5 cm., upper
base truncate more or less parallel to the rhachis, lower side cut down to the costa to a
third to half or more the length of the pinna, apex blunt or acute, upper margin and
distal part of lower margin dentate ; veins mostly once forked, continuous to the
apex of the teeth ; texture thin, herbaceous. Sori occupying the middle part of the
veins and on the acroscopic fork, sometimes confined to the distal half of the pinnae.
Spores dark, with a rather broad finely toothed perispore wing and many
anastomosing surface folds.

6a. Asplenium unilaterale var. unilaterale.

Pinnae small, 2-4 x 0-5 cm. ora little larger.

CEYLON : Thwaites C. P. 427 (BM ; K ; PDA—data on PDA sheet : Haputelle,
Apr. 1846). Thwaites C.P. 1337 (BM in part ; E in part ; GH). Thwaites C.P.
1338 (PDA—localities cited: Matale, 1847, Gardner; Palagalla, Oct. 1853 ;
Hantane, Jan. 1854). Parawatta, Matale, May 1858, Herb. Brodie (E). Deni-
yaya, 550 m., 5 Feb. 1954, Schmid 1148 (BM). Nuwara Eliya, 1,950 m., 24-27 Feb.
1954, Schmid 1352 (BM). Hakgala, by stream in jungle, 1,650 m., 27 Dec. 1950,
Sledge 741 (BM). Ramboda Pass, in jungle by track to Maturata, 1,890 m., 17 Mar.
1954, Sledge 1300 (BM; US). Ferguson (GH). Gardner 1075 in part (E).
Gardner 1077 (BM; E; K). Gardner 1338 in part (K). Hancock 52 in part
(US 1277202). 28 Mar. 1819, Moon 590 (BM). Wall (GH).

6b. Asplenium unilaterale var. majus (C. Chr.) Sledge, stat. nov.
Asplenium excisum C. Presl, Epim. Bot. : 74 (1851).—Holtt., Fl. Malaya 2: 439 (1954).
Asplenium unilaterale forma majus C. Chr. in Bernice P. Bishop Mus. Bull. 177 : 67 (1943).

Pinnae large, 5—7 X I-15 cm. or still larger.

CEYLON : Thwaites C.P. 1337 (BM in part ; K ; PDA). Corbet’s Gap, 1,200 m.,
7 Jan. 1951, Sledge 853 (BM; K). Brae, Central Province, in jungle, 1,080 m.,
4 Mar. 1954, Sledge 1238 (BM). Naylor Beckett 24 (GH). Ferguson (US 815498 in
part). Wall in Herb. Gamble (K).

On wet rocks by streams and on wet banks ; common in forests, 500—2,000 m.

Tropical Africa from Ghana, Fernando Po and Cameroons eastwards to Madagascar,
Mascarenes, Seychelles, Ceylon, North and South India, Thailand, Indochina,
Malaya, Indonesia and New Guinea, northwards to China and Japan, Philippines,
Polynesia and Hawaii.

The two extremes of size look very different. The smaller one matches Lamarck’s
A. unilaterale, the type sheet of which carries two fronds, in one of which the largest
pinnae are 2:5 cm. and in the other 4-5 cm. long. Larger plants match Cuming Ir10
from Luzon, the type of A. exciswm C. Presl, in which the largest pinnae are 6 X I°5
cm. in the Kew specimen and up to 10 cm. long in the British Museum specimen.
Some North Indian specimens have pinnae reaching 10 cm. in length.

THE-CEYLON SPECIES OF ASPLENIUM 247

Copeland (Bernice P. Bishop Mus. Bull. 59 : 61 (1929)) treated A. excisum as
merely a large form of A. wnilaterale ; and Christensen (Bernice P. Bishop Mus.
Bull. 177: 67 (1943)) recognized two forms of A. unilaterale, equating A. excisum with
his forma majus and stating that ‘‘ intermediate forms are found in Samoa as well as
in the wide area where this species occurs’. Holttum (Fl. Malaya 2: 439 (1954))
gives specific rank to A. exciswm whilst admitting it to be “ very closely related to
A. unilaterale’’, the typical form of which occurs in the same locality in Malaya as
A. excisum. He also refers to some large specimens growing amongst smaller ones
of A. unilaterale ‘‘ which seem intermediate between A. unilaterale and A. excisum”’.
Beddome’s illustration (Ferns S. Ind.: t. 132 (1864)) represents such an intermediate
form; the largest pinnae on the specimen are 5 X Icm. Several of the sori on this
gathering are allantodioid as correctly depicted in the plate.

I can find no significant difference between small and large plants and as both
appear to be coexistent throughout the range of A. unzlaterale I prefer to regard them
as variants of one species. Wall (Cat. Ferns Indig. Ceyl. : Table, 4 (1873)) also
states that the extreme forms are connected by intermediates and that the large
forms grow in “‘ wet swampy spots’. I cannot confirm that size difference is simply
an expression of habitat difference but a Ceylon plant with large pinnae produced
still larger ones (7-8 cm. long) in cultivation at Kew.

7. Asplenium obscurum Blume, Enum. Pl. Jav. 2: 181 (1828).
Asplenium cristatum Wall., Numer. List : 9, n. 211 (1829), nom. nud. ; non A. cristatum
Lam. (1786).
Asplenium erosodentatum Blume, tom. cit. : 182 (1828).
Asplenium serriforme Mett. in Abhandl. Senckenb. Naturforsch. Ges. 3: 163, t. 4 fig. 13
(1859) (“‘ serraeforme ’’).

Rhizome creeping. Stipes to 20 cm. long, grey-green ; rhachis of the same colour.
Lamina simply pinnate, 30-40 X 7-10 cm., oblong or oblong-lanceolate, with 25-35
pairs of shortly stalked spreading, falciform and dimidiate pinnae ; pinnae 4-6 x
I cm., upper base broadly cuneate to truncate, lower base cut down to the costa to
nearly half the length of the pinna, apex acute, margins serrate-dentate, often doubly
serrate above ; veins mostly once forked, continuous to the apex of the teeth ;
texture herbaceous. Sori short, about equidistant from costa and margin. Spores
dark, with a broad perispore wing with toothed edge and with anastomosing surface
folds.

CEYLON : Thwaites C.P. 3269 (BM; K). Wall (GL).

In wet mountain forests ; not common.

Malawi, Madagascar, Ceylon, South India (Nilgiri and Palni Hills), North
India, South China, Burma, Thailand, Indochina, Java.

The dull greyish-green stipe and rhachis serve to separate this readily from A.
untlaterale var. majus, in which the stipe and rhachis are dark-chestnut to black and
glossy, though in the latter the narrow green wings which extend downwards from
the pinnae on either side of the grooved upper surface of the rhachis may give a
green appearance when viewed above to a rhachis which in the same frond is dark
and glossy when seen from below. In other respects A. obscurum differs little from

248 THE CEYLON SPECIES OF ASPLENIUM

A. unilaterale var. majus, but the latter seems to be at one end of a range of forms
connecting it with A. unilaterale var. unilaterale, in which the pinnae are small. A.
obscurum apparently shows no such range of variation. Two of Wall’s specimens of A.
obscurum in Herb. Glasgow University have many completely allantodioid sori with
indusia fixed on both sides of the sori and bursting dorsally. As noted above, the
same condition has been observed in A. wnilaterale.

Christensen was the first to record this species from Madagascar, whence he cites
specimens which “ agree perfectly with Javanese ones ’’, and the Ceylon specimens
quoted above agree with those so named by Christensen.

8. Asplenium erectum Bory ex Willd. in L., Sp. Pl., ed. 4, 5: 328 (1810).—
Tardieu-Blot in Humbert, Fl. Madag., Polypod. 1: 219, fig. 30. (1-3) (1958).
(Plate 20.)

Asplenium pyramidatum Desv. in Mém. Soc. Linn. Par. 6: 271 (1827).

Asplenium camptorhachis Kunze in Linnaea 24 : 262 (1851).—Hook., Sp. Fil. 3 : 125 (1860).

Asplenium lunulatum var. sphenolobium Kunze, tom. cit. : 264 (1851).—T. Moore, Index
Fil. : 142 (1859).

Asplenium sphenolobium Zenker ex Kunze, loc. cit. (1851), nom. syn.

Asplenium brasiliense sensu Bedd., Ferns S. Ind. : 45, t. 135 (1864) ; non Raddi.

Asplenium lunulatum sensu Baker in Hook. & Baker, Synops. Fil., ed. 2 : 202 (1874) pro
parte ; non Swartz.

Asplenium lunulatum var. camptorhachis (Kunze) Bedd., Handb. Ferns Brit. Ind. : 148
(1883).

Asplenium minus sensu Alderw. van Rosenb., Malayan Ferns : 451 (1908) ; non Blume.—
Backer & Posth., Varenfl. Jav. : 135 (1939).

Asplenium sphenolobium (Kunze) Hieron. in Wiss. Ergebn. Deutsch. Z.-Afr.-Exped. 2:
14 (1910) ; in Hedwigia 60 : 226 (1919).

Asplenium lunulatum var. evectum (Bory ex Willd.) Sim, Ferns S. Afr., ed. 2 : 145 (1915).

Asplenium macraei forma sphenolobium (Kunze) Skottsb. in Medd. Géteb. Bot. Tradg. 15 :
67 (1942).

Rhizome erect, scaly at the apex, scales narrow, acute, dark-brown. Fronds
tufted, 15-50 cm. long. Stipes 1-10 cm., glabrous or nearly so, grey or dark-coloured,
not glossy ; rhachis naked, grooved above with a very narrow green wing on each side
of the groove. Lamina 10-40 X 2-4 cm., linear-elliptic with 25-40 pairs of sub-
sessile, patent pinnae, the lower and upper ones gradually reduced in size and the
lower ones commonly deflexed with the lowermost often reduced to auricles ; pinnae
I-2 X 0-5 cm., trapezoid-lanceolate, upper base truncate, more or less auricled, lower
base exciso-cuneate, the edges bluntly inciso-crenate throughout, lowermost pinnae
sometimes auricled on both bases ; basal acroscopic veins forked, the rest simple ;
texture herbaceous. Sori short, oblong, not reaching the margin or costa, usually
absent from the auricles. Spores plano-convex, 24-30 xX 18-2Iy, with an undulate
perispore wing and irregular surface folds.

CEYLON : Thwaites C.P. 3141 (BM; E; K ; PDA —localities cited on PDA
sheets : Hewahetta, Oct. 1852 ; El. Plains, Apr. 1854 ; Central Province, Mar. 1866).
Thwaites C.P. 1336 (GH). Kotmalie, 1847, Fortescue (CGE). Central Province, forests,
Naylor Beckett 191 (BM; E; GH; US 71779). Oodawella near Kandy, in jungle, 1,200

THE CEYLON SPECIES OF ASPLENIUM 249

m., 8 Dec. 1950, Sledge 531 (BM). Adam’s Peak, on rocks, 1,350 m., 14 Feb. 1908,
Matthew (K). Same locality, terrestrial in jungle, 1,950 m., 14 Dec. 1950, Sledge
626 (BM). Hakgala, jungle below the road, 22 May tg11, J. M. Silva (PDA).
Same locality, terrestrial in jungle, 1,800 m., 16 Dec. 1950, Sledge 642 (BM). Ascent
of Mt. Pedrotalagalla, in shady ground by side of track, 2,025 m., 26 Dec. 1950,
Sledge 727 (BM). Same locality and date, Holttum 39181 (SING). Nuwara Eliya,
on dry bank and on trees in forests, Sept. 1844, Gardner 1074 (CGE ; K). Same
locality, 1830, Walker (G). Same locality, 1,800 m., Apr. 1899, Gamble 27575 (K).
Same locality, in woods, 1,800 m., 10 May 1906, Matthew (K). Same locality, Mrs.
Chevalier (BM). Same locality, Freeman 131 A, 132 B, 133 C (BM). Ramboda Pass,
in forest by the Maturata track, 1,890 m., 17 Mar. 1954, Sledge 1353 (BM). Tona-
combe Estate, Namunukula, Uva Province, 1,275 m., 21 Feb. 1954, Sledge 1161
(BM). Ferguson (GH ; US 815481, 815482, 815498 in part). Hancock 33 (US
1277191). 21 Mar. 1819, Moon 483 (BM). Palliser (US 684013). Walker (K).
Wall (E; GH; GL; PDA; US 418676, 418678).

Frequent in forests in the mountains above 1,200 m.

Tropical and South Africa, Mascarene Islands, Ceylon, South India, Java, Timor,
Samoa, Hawaii.

Earlier writers followed Baker (in Hooker & Baker’s Synopsis Filicum) in using
the name Asplenium lunulatum for Ceylon plants. Swartz first used this name
(Synops. Fil. : 80 (1806)) to replace the illegitimate name A. falcatum Thunb. (non
Lam.), which was based on material from the Cape of Good Hope. But as Hooker
long ago pointed out (Sp. Fil. 3 : 128 (1860)) the plant was unknown to Swartz
and “no authentic specimen nor any authentic intelligible description exists ”’.
As currently interpreted, A. lunulatum (sensu stricto) is considered to be an
exclusively South African plant.

Several Nilgiri gatherings made by Schmid were named A. lunulatum var. spheno-
lobium by Kunze, the varietal epithet having been previously used by Zenker as
the epithet of a manuscript specific name. In the same work Kunze described A.
camptorhachis from another Nilgiri gathering, Schmid 123. The former name was
later reduced to synonymy and the latter treated as a form or (e.g. in Beddome’s
Handbook to the Ferns of British India) a variety of A. lunulatum until Hieronymus
(Hedwigia 60 : 210-266 (1919) ; op. cit. 61 : 4-39 (1919)) segregated a number of
species previously referred to A. lunulatum and A. erectum. He used the name A.
sphenolobium for Ceylon plants, specimens from Java, Samoa and Hawaii being
referred to the same species. When first publishing the name A. sphenolobium in
1910, Hieronymus attributed it to “ Zenker apud Kunze ’”’, but it had previously
only been published in synonymy and his was its first valid publication as a specific
name. Hieronymus considered that A. camptorhachis was probably a mountain
form or variety of A. sphenolobium.

I have examined Schmid 96, cited by Kunze as A. lunulatum var. sphenolobium,
and Schmid 123, the type of A. camptorhachis, both in the Botanisches Museum,
Berlin, and I agree with Hieronymus that the former is identical with Ceylon plants.
The sheet of Schmid 123 carries a single small specimen, which incidentally shows no

250 THE CEYLON SPECIES OF ASPLENIUM

‘

sign of buds on the rhachis though Kunze described it as ‘“‘ saepe gemmifera ”’.
This sheet has been annotated by Hieronymus “A. sphenolobium var. camptorhachis
(Kze.) Hieron. ”’ and I agree with him in referring the specimen to A. sphenolobium,
though in so plastic a species I do not consider the differences merit even varietal
distinction. The specimens quoted by Hieronymus from Java, Samoa and Hawaii,
and others from the same regions, are inseparable from South Indian and Ceylon
plants.

Skottsberg (Medd. Géteb. Bot. Tradg. 15 : 59-67 (1942)) has discussed the
taxonomy of A. lunulatum and its allies at length. Many names are involved but
probably few valid taxa. Large suites of specimens from Ceylon display consider-
ably greater variation in pinna form than Hieronymus or Skottsberg allow but I am
convinced that all belong to a single variable species. Sometimes the auricle is free
almost or quite to the base, when the pinna agrees with Skottsberg’s illustration
(tom. cit. : 66, fig. 87) of A. macraei forma subbipinnatum (Hilleb.) Skottsb., or with
Tardieu-Blot’s illustration (in Humbert, Fl. Madag., Polypod. 1: 221, fig. 30 (6)
(1958)) of A. evectum var. zeyheri' ; but all pinnae on the same rootstock are not
uniform and I do not think these variations are of any real taxonomic significance.
Specimens from tropical Africa named A. lunulatum var. serratodentatum Rosen-
stock and A. guintasit Gandog. I am unable to distinguish from Ceylon plants, and
Skottsberg (tom. cit. : 67) also says of South Indian plants that “‘ much the same
form is spread over Tropical Africa ”’.

It is quite evident from Skottsberg’s thorough and critical comparisons and his
series of drawings of scales, spores and pinnae of plants from Hawaii, South India,
Ceylon and Africa, that any taxonomic categories proposed within such a series
must have purely arbitrary boundaries. His discussion fully supports his conclusion
(tom. cit. : 65) that “ I cannot give these segregates a higher rank than forma. All
are connected by intermediate specimens and to some extent the shape and cutting
of corresponding pinnae varies in the same plant ’’. Christensen (Bernice P. Bishop
Mus. Bull. 25 : 26 (1925)) had earlier stated that he was “ nearly convinced ”’ that
A. macraet (which Hieronymus had treated as a distinct species) and A. spbhenolobium
were ‘‘ forms of a single species’’ and, as a result of his detailed comparisons,
Skottsberg (tom. cit. : 62) concludes “ I cannot see how a line can be drawn between
Hawaiian sphenolobium and macraei’’. He therefore reduces the former to a forma
of the latter, regretting that the rules of priority require the adoption of a specific
epithet originally given to the most extreme bipinnate form of the species, which
appears to be confined to Hawaii, whilst the commoner, simply pinnate form with
dentate or lobed pinnae ranges from there to tropical Africa.

Skottsberg, however, has little to say about A. erectum (sensu stricto). The name
is constantly coupled with A. /unulatum in his discussion and he did not examine the
type or authentic examples. <A. erectwm was described from specimens collected by
Bory in Bourbon (i.e. Réunion) and was later regarded as either a synonym or variety
of A. lunulatum until Hieronymus reinstated it. Hieronymus says it is closer to

1 Tardieu-Blot attributes this name to “ (Pappe & Rawson) Alston & Schelpe in Journ. S. Afr. bot.,
XVIII (1952), 161 ”’ but the combination was actually made first by T. Moore (Index Fil.: 127 (1859)).

THE CEYLON SPECIES OF ASPLENIUM 251

A. sphenolobium than to A. lunulatum, differing in its longer scales with different
cell net, in its wider fronds and in the more deeply serrate pinnae. As regards the
longer scales—up to 5 mm. in A. evectum—Skottsberg points out that Hieronymus’s
A. sphenolobium var. diplaziosorum has scales 5 mm. long, so the distinction can
scarcely rank as of specific value ; and one has only to look at Skottsberg’s series of
illustrations (tom. cit. : 61, figs. 36-61) of rhizome scales from plants ranging from
Hawaii to Africa to see how impracticable it is to draw even an approximate
boundary between the examples figured on the basis of size, cell shape or wall
thickness ; indeed Hieronymus himself admits (Hedwigia 60: 227) that the scales
vary in this respect in Indian plants, whilst Skottsberg states that ‘‘ the thickness
of the cell walls may vary even in the same specimen’’. As regards the reputedly
wider fronds and different depth of serration of the pinnae in A. erectum, I find no
appreciable difference in either respect between the type specimen of A. evectum and
other Réunion specimens at Kew and many examples of Ceylon and Indian plants.
One of the characters given in Willdenow’s original description was the presence of
auricles on both sides of the lower pinnae in A. erectwm and this is evident in the
type specimen. But in Réunion plants at Kew, as stated by Hooker, only “‘ some
specimens ’’ show an auricle both above and below ; moreover a frond of the type
specimen of A. pyramuidatum Desv. (equated by Hieronymus and Christensen with
A. erectum) is without auricles on the lower sides of the basal pinnae. No valid
distinction remains therefore whereby A. evectum may be distinguished from
A. sphenolobium.

Christensen was evidently not convinced of the soundness of Hieronymus’s
conclusions for in his Pteridophytes of Madagascar (1932) he did not follow him in
the treatment of A. lunulatum (sensu lato) but adopted Sim’s arrangement in placing
A. erectum as a variety under A. lunulatum, the form with auricles on both sides of
the pinna being described as A. lunulatum var. erectum subvar. subbiauritum. Later
(Index Fil., Suppl. 3: 32 (1934)) he cited A. evectum as a South African fern without
mention of the Mascarene Islands whence the type was described, and in his Pteri-
dophytes of Tristan da Cunha (Res. Norw. Sci. Exped. Tristan da Cunha 1 (6): 13
(1940)) he gave specific rank to A. erectwm and so identified plants from the Tristan
da Cunha group of islands as he could “ not find any good character by which most
specimens from the islands may be distinguished from A. erectum’’.

The type specimen of A. evectum is in the Willdenow Herbarium at Berlin. There
is a photograph of it in the British Museum Herbarium (see Plate 20) and two sheets
from “ Herb. Mus. Paris. Ile Bourbon” in Hooker’s collection at Kew. Though
neither of these sheets has the collector’s name inscribed on the label, the specimens
on one or both sheets are presumably from Bory’s collection as in Hooker’s citation of
specimens (Sp. Fil. 3: 127 (1860)) he says “I... confine myself to specimens in my
own herbarium’, the first citation being “ Bourbon Bory. Herb. Mus. Paris in
Herb. Nostr.”” These and other examples from Réunion and the other Mascarene
Islands are, to me, indistinguishable from Ceylon plants.

It has always been recognized that the South African plant regarded as the true
A. lunulatum is very close to A. erectum, but many South African specimens named

252 THE CEYLON SPECIES OF ASPLENIUM

A. lunulatum have considerably less incised pinnae and the illustration in Sim (Ferns
S. Afr., ed. 2: t. 47 A, B (1915)) of A. lunulatum var. lunulatum shows a frond with
pinnae much more nearly entire than in Ceylon and South Indian plants. I do not
know if there are consistent differences in scale and spore characters between African
plants corresponding with Sim’s A. lunulatum var. lunulatum and Asiatic plants, but
as the former are similar to, rather than identical with, Ceylon and Indian plants I
am not disposed to unite them. I conclude therefore that Aspleniwm erectum is the
correct name for Ceylon plants.

g. Asplenium inaequilaterale Willd. in L., Sp. Pl., ed. 4, 5: 322 (1810).—
Hieron. in Hedwigia 61 : 22 (1919).
Asplenium brachyotus Kunze in Linnaea 10 : 512 (1836).
Asplenium trapeziforme sensu Bedd., Ferns S. Ind. : 45, t. 134 (1864) ; non Roxb.
Asplenium lunulatum var. trapeziforme Bedd., Handb. Ferns Brit. Ind. : 148 (1883) pro
parte ; non A. trapeziforme Roxb.

Rhizome erect, scaly at the apex, scales linear, tapering, entire, dark-brown-black,
3-5 mm. long. Fronds tufted, 20-40 cm. long, rarely more. Stipes 8-16 cm. long,
grey, sparsely and minutely hairy. Lamina up to 25 cm. long, 4-8 cm. broad,
oblong-lanceolate, terminating in a lobed apex, simply pinnate with 15-20 pairs of
shortly stalked pinnae, the lowermost not reduced ; pinnae 2:5-5 x Icm., trapezoid-
lanceolate, dimidiate, upper base truncate and more or less parallel to the rhachis,
lower margin obliquely cut away to 4 or more of the length of the pinna, apex acute
or subacute, both margins crenate-dentate, glabrous above and beneath ; veins
forked, not continuous to the apex of the teeth ; texture herbaceous. Sori on the
anterior forks of the veins forming a row on each side of the costa but not touching
the costa and falling well short of the margin. Spores plano-convex, 30-33 X 2I-24n,
with a perispore forming a median wing and anastomosing surface folds.

CEYLON: Thwaites C.P. 1336 (BM; GH; PDA—data on PDA sheet:
Hantane, July 1847, Gardner ; Pallagalla, Oct. 1853). Thwaites C.P. 1337 in part (E).
Hantane range, in forests, Oct. 1844, Gardner 1075 (CGE). Hakgala, common
in jungle, cult. in R.B.G., Senaratna 805 (PDA). Kandy, Mrs. Chevalier (BM).
Same locality, Freeman 127 A, 128 B (BM). Labukellia, Freeman 129 c (BM).
Ramboda Pass, Freeman 130 D (BM). Corbet’s Gap, I,200 m., 7 Jan. 1951,
Sledge 855 (BM). Same locality, 22 Jan. 1954, Sledge 1033 (BM; K). MHunnas-
giriva, Central Province, by stream in jungle, 870 m., 16 Jan. 1954, Sledge 980
(BM ; K). Brae, Central Province, in wet ground in forest, 1,080 m., 4 Mar. 1954,
Sledge 1239 (BM; K; US). Naylor Beckett 2480 (GH). Ferguson (GH; US 815510).
1847, Gardner 29 in Herb. Boissier (G). Gardner 1075 in part (E). Gardner 1336
(K). Gardner 1338 in part (K). Hancock 52 in part (US 1277202). Robinson C56
(K). 1853-55, Thwaites in Herb. Boissier (G). Wall (GL; PDA; US 1095878).

On damp ground in mountain forests, 600—1,900 m.

South America, tropical and South Africa, Madagascar, Réunion, Ceylon, South
India (Nilgiri, Palni and Anaimalai Hills, Bombay).

A. maequilaterale has been confused with A. unilaterale and A. obscurum to which
its fronds bear the closest resemblance. It differs from both these species in its erect

THE CEYLON SPECIES OF ASPLENIUM 253

rhizomes bearing tufted fronds, the stipes and rhachides of which are not, as in A.
unilaterale, dark-purple-black and polished. In A. inaequilaterale no separate
pinnae are formed in the uppermost part of the frond, which terminates in a lobed
apex; whereas in A. unilaterale and A. obscurum the distal pinnae decrease regularly
in size but remain discrete to the apex of the frond. In A. imaequilaterale the lamina
on the lower margins of the pinnae is obliquely cuneate, meeting the costa near its
base, whereas in A. unilaterale and A. obscurum the lower margins of the pinnae are
cut down to the costa for a considerable distance from the base. The lateral veins of
the pinnae run to the tips of the marginal teeth in A. wnilaterale and A. obscurum ;
in A. inaequilaterale the veins fall short of the margin.

Specimens from Réunion, whence the type was described, are a good match for
Ceylon and South Indian plants.

Roxburgh’s A. trapeziforme, described from the “ Malay Islands ’’, is not the
same plant. It also has an erect rhizome but photographs in the British Museum
Herbarium of the type specimen, including enlargements of the pinnae, clearly show
the veins running to the extremities of the marginal teeth. The fronds are also
pinnate to the end, not terminating in a distal, lobed, pinna-like extremity.

10. Asplenium tenerum Forst. f., Florul. Ins. Austr. Prodr. : 80 (1786).
Asplenium elongatum Swartz, Synops. Fil. : 79 (1806).—Hook., Sp. Fil 3 : 117 (1860).—
Thw., Enum. Pl. Zeyl. : 383 (1864).
Ashplenium tenerum var. terminans Mett. in Abhandl. Senckenb. Naturforsch. Ges. 3 : 113
(1859).

Rhizome erect, clothed at the apex with lanceolate, acute, dark-coloured scales.
Stipes tufted, green, up to 20 cm. long, scaly when young, becoming glabrescent.
Lamina 15-30 X 4-8 cm., oblong-lanceolate, simply pinnate with 10-25 pairs of
spreading pinnae, the lower ones not or scarcely reduced, the upper ones suddenly
reduced, the frond terminating in a narrow, winged prolongation, 2-3 cm. long, bearing
simple oblique lobes ; pinnae 2-5 cm. long, 6-15 mm. broad, shortly stalked, oblong-
lanceolate, upper base more or less truncate, often subauriculate, lower base narrowly
cuneate, margins with regular, oblique, blunt teeth, apex usually bluntly rounded,
sometimes acute, both surfaces glabrous or with a few minute, appressed, pale-brown,
irregular scales on the veins beneath ; costa raised above, veins all simple save the
basal one on the acroscopic side which is forked once or twice ; texture firm-herbaceous.
Sori not reaching either the costa or the margin ; indusium pale. Spores light-
brown, plano-convex, 42-48 X 27-33y, with a winged perispore and a few surface
folds.

CEYLON: Thwaites C.P. 1007 (BM; E; GH; K; PDA—data on PDA sheets:
Maturata, 1819, Moon ; Aliagalla, Pelagalla, Oct. 1853). Poorkpittia Pass, Matale,
Dec. 1860, Sept. 1864, Herb. Brodie (E). Peacock Hill, Pussalawa, Robinson (K). Same
locality, 1870-71, Randall in Rawson 3220/297 (BM). Summit of Ritigala, North
Central Province, 24 Mar. 1905, Willis (PDA). Kabot’s Gap, Rangala, 8 Sept. 1927,
Alston (PDA). Oodawella near Kandy, in jungle, 1,200 m., 8 Dec. 1950, Sledge 530
(BM). Corbet’s Gap, in shady forest, 1,200 m., 7 Jan. 1951, Sledge 851 (BM). Naylor

254 THE CEYLON SPECIES OF ASPLENIUM

Beckett 23 (GH). Ferguson (GH; PDA; US 815516). Gardner 26 (BM). Gardner
1078 (BM; E; K). Hancock 29 (US 1277188). 4 Feb. 1819, Moon roqg (BM).
Robinson 53 (K). Walker (K). Wall (E ; GL; K).

In mountain forests, I,200—2,100 m.

Seychelles, Ceylon, South India, Burma, Thailand, Indochina, Malaya, Java,
Borneo, Philippines, New Guinea, Polynesia.

Ceylon plants agree closely with examples from Polynesia, whence the type was
described. Mettenius’s variety ¢eryminans was described from material of Gardner’s
from Ceylon, said to differ in its more coriaceous fronds, which were paler beneath,
and in its longer pinnae. It was evidently founded on a single gathering and
represents no more than a minor state or form. Iam unable to see any significant
difference between Gardner’s specimens at Kew or other Ceylon plants and those
from elsewhere.

11. Asplenium pellucidum Lam. in Encycl. Méth., Bot. 2 : 305 (1786).
Ashlenium hirtum Kaulf., Enum. Fil.: 169 (1824).—Hook., Sp. Fil. 3: 149 (1860).—
Bedd., Handb. Ferns Brit. Ind. : 149 (1883).

Rhizome short-creeping, clothed with subulate, black, hair-like scales. Fronds
60-90 cm. long, 10 cm. or more broad. Stipes and rhachides dark, grooved above,
fibrillose with black, hair-like scales. Lamina simply pinnate, narrowed above and
below with 60 or more pairs of pinnae ; pinnae horizontal, the lower ones gradually
reduced to short auricle-like appendages, sessile, up to 6 x I cm., linear-oblong,
apex acute, broadly cuneate or truncate and auricled at the upper base, narrowly
cuneate at the lower base, margins serrate ; veins once or twice forked, the costa
grooved above, hairy below when young ; texture chartaceous to subcoriaceous,
dark-brown when dry. Sori on the acroscopic branches of the veins forming two
oblique rows extending from the costa and falling short of the margin ; indusium
firm, brown.

CEYLON: Near Galle (PDA).

Habitat in Ceylon unknown.

Madagascar, Mascarene Islands, Ceylon, Sikkim, Burma, Indochina, Malaya,
Sumatra, Java, Borneo, Philippines, New Guinea.

The distribution of A. pellucidum is consistent with its occurrence in Ceylon but
the single frond in the Peradeniya collection, added since Thwaites’s time, is the
only specimen I have seen.

12. Asplenium zenkeranum Kunze in Linnaea 24: 259 (1851) (“ Zenkerianum”’”).
—Bedd., Handb. Ferns Brit. Ind. : 148 (1883).
Asplenium persicifolium var. latifolium Hook., Sp. Fil. 3 : 109 (1860).—Thw., Enum. PI.
Zeyl. : 383 (1864).

Rhizome subrepent, clothed with lanceolate, acute, brown scales with paler
fimbriate margins. Stipes grey-green, up to 30 cm. long, grooved above and bearing
narrow, brown, fimbriate scales ; rhachis similarly grooved and scaly. Lamina
oblong-lanceolate, 30-70 X 10-20 cm., imparipinnate with 7-20 pairs of distant

THE CEYLON SPECIES OF ASPLENIUM 255

pinnae and a terminal pinna bearing a scaly bud on the costa ; pinnae normally
5-I0 X I°5-2:5 cm., sometimes to 15 X 3 cm., stalked, lanceolate, oblong-lanceolate
or ligulate, apex acute, upper base rounded, lower base cuneate, margins crenate
or subentire, costa raised above, upper surface naked, lower surface usually with a
few appressed, laciniate, ferrugineous scales ; veins once or twice forked ; texture
firm, chartaceous. Sori forming two oblique rows between the costa and the margin;
indusium pale, broad. Spores plano-convex to ellipsoid, 42-50 x 30-40p, with a
rather broad undulate perispore wing and a few surface folds.

CEYLON: Thwaites C.P. 3461 (BM ; E; GH; K; PDA). Central Province,
goo-1,200 m., Naylor Beckett 1123 (E ; GH). Corbet’s Gap, in shady forest, 1,200 m.
7 Jan. 1951, Sledge 833 (BM ; US). Hakgala, on rocks by stream in ravine, 1,650 m.,
26 Feb. 1954, Sledge 1214 (BM). Same locality, terrestrial in forest, 1,650 m.,
20 Mar. 1954, Sledge 1339 (BM). Nuwara Eliya, Freeman 197 A (BM). Same locality,
2 Sept. 1926 (PDA). Moon Plains jungle, Freeman 198 B (BM). 1899, Bradford
470 (K). Ferguson (GH; US 815521). Gardner (K). Robinson 51 (K). Walker
(K). Wall (BM; E; GL; K).

In moist forests of the Central Province, I,200-2,100 m.

Ceylon, South India (Nilgiri and Palni Hills).

The proliferous terminal pinna in this species is usually contracted suddenly to
the costa at the point where the large scaly bud is formed, and the costa is then
prolonged and develops an irregular lamina. Very occasionally fronds may continue
apical growth more or less indefinitely without forming a terminal pinna.

13. Asplenium serricula Fée, Mém. Fam. Foug. 5 : 196 (1852).
Asplenium wightianum Wall., Numer. List : 66, n. 2215 (1830), nom. nud.
Asplenium coviaceum Bory in Bélanger, Voy. Ind.-or., Bot. 2: 46 (1833); non A.
coriaceum Desv. (1827).
Asplenium wightianum Wall. ex Hook., Sp. Fil. 3: 105, t. 167 (1860)—Bedd., Handb. Ferns
Brit. Ind. : 146 (1883).

Rhizome oblique, subrepent, scaly at the apex, scales linear, tapering, 6 x I mm.,
walls brown, margins paler, fimbriate. Fronds tufted, 25-50 cm. long. Stipes
green, up to 20 cm. long, scaly at the base, naked above. Lamina imparipinnate,
pale-glaucous-green, up to 30 X I5 cm., sometimes more, with 3-Io pinnae on
each side of the rhachis and a terminal pinna resembling the lateral ones ; pinnae
shortly stalked, linear or elliptic-oblong, 7-15 cm. long, about 1 cm. (rarely 2 cm.)
broad, widest at or near the middle, narrowing gradually to the base, apex long-acuminate,
margins coarsely serrate, glabrous on both surfaces or with a few laciniate pale-
brown scales beneath, costa raised above ; veins simple or once forked ; texture
coriaceous or subcoriaceous. Sori 0:5-I cm. long, extending from near the costa
but falling short of the margin ; indusium pale, whitish. Spores reniform to
plano-convex, 42-48 X 27—30u, with a winged perispore and surface folds.

CEYLON : Thwaites C.P. roro in part (BM; E; G; K ; PDA—data pencilled
on PDA sheets: Nuwara Eliya, 1847 and Feb. 1857, Gardner ; Matale, 1849,
Gardner). Hantane Range, on trees in forest, Oct. 1844, Gardner 1070 bis (CGE).

256 THE CEYLON SPECIES OF ASPLENIUM

Oodawella, Robinson 69 (K). Summit of Ritigala, 24 Mar. 1905, Willis (PDA).
Central Province, in forest, Wall in Herb. Hance (BM). Cabragalla, Matale, June
1858, Herb. Brodie (E). Rattotte, Matale, 21 Dec. 1862, Herb. Brodie (E).
Hunnasgiriya, on bank of stream in jungle, 870 m., 16 Jan. 1954, Sledge 979 (BM).
Brae, in jungle by track to Hoolankande Pass, 1,080 m., 4 Mar. 1954, Sledge
1237 (BM). Naylor Beckett 212 (GH). Ferguson (GH ; US 815522). Gardner
zo7o in part (BM; E; K). Cult. Colombo, Apr. 1899, Gamble 27541 (K).
Bradford in Herb. Hance (BM). Hancock 43 (US 1277198). Levinge in Herb.
Botssier (G). 17 Feb. 1819, Moon 223 (BM). Walker ex Herb. Mus. Paris. 521
(i); “Wall (BM3 Es -GH 3s GL, Kk).

On trees and rocks in mountain forests ; infrequent.

Ceylon, South India, Philippines.

Fée described this species from Ceylon specimens—‘‘ Gardner no. 30 ”—and his
reference to the rigid texture and pallid green colour clearly indicate that his plant
was the same as that which Hooker later described as A. wightianum. Its differences
from A. longipes are referred to under that species. South Indian specimens are
identical with those from Ceylon. It appears to be more widely distributed in
Ceylon than A. longipes but at lower elevations.

A Philippine specimen at Kew seems identical with Ceylon plants. Copeland
included A. serricula (as A. wightianum) in his Polypodiaceae of the Philippine
Islands (1905) but he omits it in his Fern Flora of the Philippines (1960).

14. Asplenium longipes Fée, Mém. Fam. Foug. 5 : 195 (1852) ; op. cit. 7: 49,
t. 16 fig. 3 (1857).
A shplenium walkerae Hook., Sp. Fil. 3 : 108, t. 163 (1860).
Asplenium vulcanicum sensu T. Moore, Index Fil. : 178 (1860) pro parte, quoad specim.
Zeyl. ; non Blume.—Hook. & Baker, Synops. Fil. : 201 (1867) pro parte.

Rhizome obliquely erect, scaly at the apex, scales tapering, 7-8 xX I-I'5 mm.,
walls brown, margins paler, fimbriate. Fronds tufted, 30-60 cm. or more long.
Stipes green, up to 30 cm. long, scaly when young, becoming naked or almost so
with age. Lamina imparipinnate, green, up to 40 X 15 cm., with 4-8 pinnae on
each side of the rhachis and a terminal pinna resembling the lateral ones ; pinnae
petiolate, lanceolate, 7-12 cm. long, about 2-2-5 cm. (rarely 3:5 cm.) broad, widest
near the base, superior base rounded to broadly cuneate, inferior base more obliquely
cuneate, apex acuminate, margins coarsely serrate, glabrous on both surfaces or
with a few small brown scales beneath, costa raised above ; veins simple or once
forked ; texture firmly herbaceous. Sori up to.1°5 cm. long, extending from the
costa to near the margin ; indusium whitish. Spores plano-convex, 36-42 xX
24-271, with a winged perispore and surface folds.

CEYLON : ThwaitesC.P. roroin part (BM ; E; G; GH; K ; PDA). Ramboda
Pass, on trees in forests, Sept. 1844, Gardner 1070 (CGE). Same locality, 1868,
Randall in Rawson 3220/56 (BM). Nuwara Eliya, 1,500-1,800 m., Wall in Herb.
Hance (BM). Same locality, Mrs. Chevalier (BM). Same locality, 1,800 m., Free-
man I50 A, I5I B, 152 C (BM). Same locality, 1,950 m., 24-27 Feb. 1954, Schmid
1762 (BM). MHakgala, jungle below the road, 22 May 1911, J. M. Silva (PDA).

THE CEYLON SPECIES OF ASPLENIUM 257

Same locality, in jungle, 1,800 m., 16 Dec. 1950, Sledge 635 (BM). Same locality, in
jungle, 1,650 m., 26 Feb. 1954, Sledge 1215 (BM). Same locality, in jungle above
Botanic Garden, 1,800 m., 23 Dec. 1950, Sledge 707 (BM). Same locality and date,
Holttum 39171 (SING). Parawella Falls, Kandapola Forest Reserve near Nuwara
Eliya, in jungle, 1,425 m., 19 Mar. 1954, Sledge 1332 (BM; US). 1899, Bradford
(K). Ferguson (GH; US 815520). Gardner ro70 in part (BM; K). Gossett (K).
Robinson 50A (K). Walker (K, type of A. walkerae). Wall (E; GH; GL; K). Wight
1958 (E).

On trees and rocks in forests ; common about Nuwara Eliya but not recorded
from elsewhere.

Endemic.

This species was described from Ceylon plants gathered by Walker. Fée’s descrip-
tion of the pinnae as ovate-lanceolate and his illustration leave no doubt that this
is the plant which Hooker redescribed, again from specimens of Walker’s, as
A. walkerae. It is closely related to A. serricula but differs in its thinner texture,
greener fronds and in its pinnae being broadest near the base with the acroscopic
margin broadly cuneate or even rounded. In A. serricula the fronds are whitish-
green or glaucous-green and subcoriaceous in texture and the pinnae are narrowly
oblong or elliptic-oblong, broadest at or near the middle and tapering at both ends.
In A. longipes the pinnae are normally 2-2-5 cm. wide in the widest part and still
wider in robust specimens, whereas in A. serricula they are normally I cm. wide,
occasionally reaching 2 cm. in large specimens. As correctly described by Hooker,
diplazioid sori sometimes occur in A. longipes.

Thwaites (Enum. Pl. Zeyl.: 383 (1864)) considered the two to be connected by
intermediate forms and hence untenable as separate species. Wall (Cat. Ferns
Indig. Ceyl.: 4 (1873)), whose field knowledge of Ceylon ferns was both extensive
and accurate, expressed precisely the opposite view. My own experience leads me
to agree with Wall ; and nearly all herbarium specimens can be readily assigned to
one or the other species. Moreover, if A. longipes were merely a variant of
A. serricula with broad pinnae it might well be expected to accompany that species
in South India, whereas all specimens from there match Ceylon examples of
A. serricula and none approaches A. longipes.

Hooker considered the Ceylon plant ‘undoubtedly quite distinct’”’ from
A. vulcanicum Blume, but Moore (Index Fil.: 178 (1860)) and Baker (Hook. &
Baker, Synops. Fil.: 201 (1867)) included it in that species. I have examined the
type specimen of A. vulcanicum from Java and concur with Hooker’s view. It
differs in the fronds being proliferous from a bud borne distally on the rhachis and
in the oblong or oblong-lanceolate, cuneate-based pinnae, the margins of which
are subentire except towards their serrate apices. In A. longipes the fronds are never
proliferous and the margins of the lanceolate pinnae are coarsely serrate throughout.
The non-proliferous A. salignum Blume of Malaya and Java is also much less coarsely
serrate and has fewer pinnae—its fronds vary from simple to pinnate with 3-4 pairs
of pinnae—than Ceylon plants. The type sheet of this has all the fronds on three
complete specimens either simple or trifoliolate. A specimen in the British Museum

258 THE CEYLON SPECIES OF ASPLENIUM

Herbarium from Java, collected by Horsfield and named A. walkerae, has coarsely
toothed pinnae as in A. longipes but differs from that in being proliferous.

A. longipes is common in the forests of the Nuwara Eliya district but I have
not met with it elsewhere. Where localities are cited on herbarium sheets, they are
all in this district.

15. Asplenium gardneri Baker in Gard. Chron. 1873: 712 (1873).—Bedd.,
Suppl. Ferns S. Ind. & Brit. Ind. : Io, t. 355 (1876).
Asplenium caudatum sensu Hook., Sp. Fil. 3 : 152 (1860) quoad specim. Zeyl. pro parte ;
non Forst. f.
Asplenium macrophyllum sensu Hook., tom. cit. : 158 (1860) pro parte ; non Swartz.—
Thw., Enum. Pl. Zeyl. : 384 (1864).
Asplenium tenerum sensu Thw., op. cit. : 383 (1864) ; non Forst. f.

Rhizome creeping, clothed at the apex with short, ovate, blackish scales 1 mm. long.
Fronds 20-50 cm. long. Stipes green, darkening with age, 5-15 cm. long, terete
on the back, channelled above, scaly at the base, elsewhere, like the rhachis, with
scattered short, broad, dark-brown scales with peltate bases and often with irregu-
larly dentate margins. Lamina oblong-lanceolate, simply pinnate, 10-30 or more
cm. long, 5-10 (15) cm. broad, green when dry, with 8-15 (20) pairs of pinnae, the
lowermost not or scarcely reduced, terminating in an apical pinna similar to the
lateral ones and gemmiparous from the upper base of one of the distal pinnae ; pinnae
3-6 (8) cm. long, 1-1-5 cm. broad near the base, shortly stalked, patent or ascending,
not imbricating, upper base broadly cuneate or truncate, lower base narrowly
cuneate, lanceolate, acute, margins toothed, glabrous above, sparsely scaly about the
base beneath and often with a few minute scales on the veins ; veins immersed,
forked, well spaced, not striate above ; texture firm-herbaceous. Sori up to I cm.
long, spreading along the veins from near the costa and falling short of the margin.
Spores plano-convex, 36-42 Xx 24-27u, with a perispore forming a rather broad
irregular undulate wing and a few surface folds.

CEYLON : Thwaites C.P. 13340 (BM ; E; GH; K; PDA). Thwaittes C.P. 1371
(K). Gardner 1072 (BM ; E ; CGE ; K—the Cambridge sheet localized as Nuwara
Eliya, on rocks in forests, June 1845). Hunnasgiriya, Beckett in Herb. Brodie
(E). Matale, Nov. 1861, Herb. Brodie (E). Deniyaya, Southern Province, in
forest above Beverley Estate, 900 m., 12 Mar. 1954, Sledge 1284 (BM). Same locality,
780 m., 4 Apr. 1954, Sledge r400 (BM; US). 1899, Anderson (E). Ferguson
(GH ; PDA; US 815491). Henderson (E). 1880, Hutchison (E). Wall (BM;
Pee GSAS):

On rocks and trees in forests, especially in the Southern Province.

Ceylon, Sumatra.

Hooker included A. gardneri partly under A. caudatum Forst. f. and partly under
A. macrophyllum Swartz. Beddome considered it ‘‘ probably only a variety ”’ of
the latter (Handb. Ferns Brit. Ind.: 151 (1883)). But it is a quite distinct species
not, in my opinion, closely related to the A. falcatum group. From the least-divided
forms of A. falcatum it differs in its creeping rhizome with much smaller and blunter
scales, in its gemmiparous fronds of a more supple and pliable texture, in the veins

THE CEYLON SPECIES OF ASPLENIUM 259

of the pinnae being immersed and not forming striations above and also being far
fewer and more widely spaced than in that species and hence in having fewer sori
to the pinna, usually five or six on each side of the costa as against three or four
times that number in A. falcatum. From the creeping A. decrescens it is at once
distinguished by its sori not forming parallel rows close to the costa and by its
gemmiparous habit.

I have only met with this species in the Sinha Raja forest and Wall cites it only
from the Southern Province. Gardner collected it at Nuwara Eliya but I have seen
no other gatherings from that much-botanized area. Teschemacher’s specimens
from Sumatra quoted by Hooker under A. macrophyllum are identical with Ceylon
plants but the Khasi Hills plants referred by C. B. Clarke to A. gardneri are
specifically distinct (A. khastanum Sledge in Kew Bull. 15 : 397 (1962)).

16. Asplenium decrescens Kunze in Linnaea 24 : 261 (1851).

Asplenium contiguum sensu T. Moore, Index Fil. : 121 (1859) quoad specim. Zeyl.; ?an
Kaulf.—Bedd., Ferns S. Ind. : 47, t. 140 (1864).

Asplenium caudatum sensu Hook., Sp. Fil. 3 : 152 (1860) quoad specim. Zeyl. pro parte ;
non Forst. f—Thw., Enum. Pl. Zeyl.: 384 (1864) pro parte.—Bedd., Handb. Ferns
Brit. Ind. : 151 (1883).

Rhizome long-creeping, clothed with narrow, finely pointed, very dark-coloured,
imbricating scales 5 mm. long. Fronds 30-60 cm. long. Stipes 5—30 cm. long, deep
purplish-brown, scaly at first, becoming almost or completely glabrous save at the
base; rhachis grooved above, usually with scattered, filiform, brown scales or more
rarely almost glabrous. Lamina oblong-lanceolate, up to 30 cm. or more long, with
10-20 pairs of shortly stalked pinnae, the lowermost slightly reduced, the uppermost
gradually reduced and merging into the lobed apex ; pinnae 5-10 cm. long and
I cm. broad near the base, lower base narrowly cuneate, upper base broadly cuneate
or rounded, narrowed gradually to the acuminate apex, both edges more or less
deeply lobed, the lobes oblique and sharply toothed, becoming simply serrate distally,
striate above and dark-olive-brown when dry, under surface with brown, filiform
scales on the veins ; veins numerous, crowded, forked ; texture chartaceous. Sov
situated on the vein branches nearest the costa, at first somewhat imbricating and
diverging but becoming confluent with age to form two parallel rows close to the costa
but with a few flabellately arranged sori on the basal lobes. Spores plano-convex,
36-42 X 24-27u, with a perispore forming an undulate wing and a few surface folds.

CEYLON : Thwaites C.P. 3140 (BM; E; GH; K; PDA). Adam’s Peak, near
summit, 2,200 m., Matthew (K). Same locality, 1,950 m., 14 Dec. 1950, Sledge 620
(BM ; US). Hulangamuwa, Matale, June 1860, Herb. Brodie (E). Matale,
Nov. 1861, Herb. Brodie (E). Jelumalai, Namunukula, 12 Mar. 1907, J. M. Silva
(PDA). Near Hakgala, terrestrial, 1,650 m., 23 Dec. 1950, Holttum 39159 (SING).
Nuwara Eliya, Freeman 139 B, 140 A, 142 C (BM). Pedrotalagalla, Nuwara Eliya,
epiphytic by path in forest, 6 Apr. 1959, Sinclair rorr7 (E). Corbet’s Gap, in second-
ary jungle, 1,320 m., 9 Dec. 1950, Sledge 568 (BM). Between Pattipola and Horton
Plains, epiphytic in forest, 1,950 m., 18 Dec. 1950, Sledge 698 (BM). Hoolankande,
Central Province, 1,350 m., 20 Jan. 1954, Sledge rorg (BM). Le Vallon, 1,500

260 THE CEYLON SPECIES OF ASPLENIUM

m., 9 Feb. 1954, Sledge rr104 (BM). Naylor Beckett 292 (GH). Ferguson (GH;
US 815479). Gardner ro8t (CGE ; K). Palliser (GH). Walker (K). Wall (E. ;
GH).

On rocks and trees in the higher parts of the interior, I,300—2,200 m.
Ceylon, South India (Nilgiri and Palni Hills).

Kunze described this species from specimens collected by Schmid in the Nilgiri
Hills. One of the two gatherings which he quotes, Herb. Koch n. 122, is in the Berlin
Botanical Museum. I have examined this sheet and have no doubt that the speci-
mens belong to the same species as the Ceylon plants which have long passed under
the name A. caudatum Forst. f. One of the specimens is complete with rhizome
and shows the creeping habit. The fronds on this plant and a separate larger frond
mounted on the same sheet show the characteristic arrangement of the sori, which
are elongated close to and more or less parallel with the costae of the pinnae. The
type specimen represents a considerably smaller form with more entire pinnae than
is commonly met with in this species, but both in size and lobing of the pinnae
A. decrescens is a variable species and the original specimens may well represent
high-altitude or precociously fertile examples. Kunze’s choice of specific epithet
was both inappropriate and inexplicable since his description states that the fronds
are not decrescent.

Hooker (Sp. Fil. 3 : 152 (1860)) was responsible for referring the Ceylon plant to
Forster’s A. caudatum. Forster (Florul. Ins. Austr. Prodr.: 80 (1786)) does not
state where his A. caudatum was collected but according to Schkuhr (Krypt. Gew.:
72, t. 77 (1809)) it came from the Friendly Islands. The type specimen at Gottingen
labelled ‘‘Asplenium caudatum Forst. prod. no. 432. In ins. Maris pacifici leg. Forster’
consists of a portion of a pinnate frond about 12 cm. long with pinnae about 4 cm.
long, pinnatifid about half-way to the costa with serrate lobes and with diverging
sori. Schkuhr’s illustration depicts the form of the pinnae and arcuate, diverging
sori, some reaching almost to the frond margin, very well. Yet Hooker (loc. cit.),
although referring to this illustration as ‘“‘ faithful’, describes the sori as “‘ almost
parallel with and near the costa’’, a statement which was no doubt responsible
for the confusion surrounding the application of this name by Beddome, Thwaites
and others who relied largely on Hooker’s work. Most specimens labelled A. cauda-
tum from south-east Asia and the Pacific Islands are easily distinguished from Ceylon
plants by the cutting of the pinnae and especially by the stipe and rhachis which are
more profusely clothed with black, clathrate scales with long hair-points.

Ceylon plants have also been referred to A. contiguum Kaulf. by Moore and
Beddome, and this identification is certainly to be preferred to Hooker’s. Kaulfuss’s
species was described from Hawaii and some specimens from there are very like
A. decrescens. Authentic specimens, however, are not available for study and
Hawaiian plants labelled A. contiguum are decidedly diverse. A Kew specimen so
identified by Christensen differs from the Ceylon plant in its less-cut pinnae which
are glabrous beneath. The Philippine A. lepturus J. Smith ex C. Presl, which
Christensen (Index Fil.: 118 (1905)) cites as a synonym of Kaulfuss’s species and to

THE CEYLON SPECIES OF ASPLENIUM 261

which Moore (Index Fil. : 121 (1859)) and Hooker (Sp. Fil. 3 : 156 (1860)) gave varietal
rank also under A. contiguwm Kaulf., is another closely related taxon. This was
based on Cuming 211 from Luzon, the specimens of which at Kew and the British
Museum differ from the Ceylon plant in their more caudate pinnae, glabrous beneath
and with more diverging sori. Ceylon and Nilgiri plants fall between these species
and the African A. friestorum C. Chr., which differs in its broader-based, thinner-
walled, brown, shining and iridescent scales, and in the position of its sori which
are all strictly costal.

Skottsberg (Medd. Géteb. Bot. Tradg. 15 : 97 (1942)) discussed the relationships
of Hawaiian, Philippine, Ceylon-South Indian and African plants and concluded
that all are conspecific with A. contiguum although he cited differences (tom. cit.: 103)
in the scale and spore characters both of the African and Ceylon-South Indian
plants and these are listed in his table (tom. cit.: 82) as varieties—unnamed—of
A.contiguum. The African A. friestorum seems to me to merit the specific separation
which all later authors have upheld and I prefer to use the name A. contiguum for
Hawaiian and Pacific Island plants and to retain Kunze’s name for Ceylon plants,
which are certainly identical with those from South India.

A. decrescens is readily distinguished from A. falcatum, which it most closely
resembles, by the position of its sori which lie close to and parallel with the costa
of the pinna, and by its long-creeping rhizome. The two species are also quite
distinct in their distribution in Ceylon, A. falcatum being a fern of low elevations
(I have not met with it above 675 m.) whereas A. decrescens grows only in the higher
mountains of the interior, above I,300 m.

17. Asplenium falcatum Lam. in Encycl. Méth., Bot. 2: 306 (1786).—Bedd.,
Handb. Ferns Brit. Ind. : 150 (1883).
Trichomanes adiantoides L., Sp. Pl. 2 : 1098 (1753).
Tarachia falcata (Lam.) C. Presl in Abhandl. K. Béhm. Ges. Wiss., Folge 5, 6 : 437 (1851).
Asplenium adiantoides (L.) C. Chr., Index Fil. : 99 (1905) ; non A. adiantoides Lam.
(1786).—Holtt., Fl. Malaya 2: 431 (1954).

Rhizome oblique or short-creeping with close-set stipes, clothed with narrow, pointed,
dark scales 5 mm. long. Fronds 30-90 cm. long. Stipes up to 30 cm. long, dark-
purple to black with spreading scales at the base and smaller, appressed, deciduous
scales with more or less stellately lobed bases above ; rhachis grooved above with
scattered, dark-brown, irregularly lobed scales. Lamina oblong-lanceolate, simply
pinnate or bipinnate, up to 40 cm. long or sometimes more, with 5-15 pairs of shortly
stalked pinnae, the lowermost not reduced, terminating in an apical pinna similar
to the lateral ones and often with one or two long ascending pinna-like lobes at its
base ; pinnae lanceolate, 5-15 cm. long, 2 cm. broad near the base, the lower base
more or less excised, the upper base broadly cuneate or rounded, narrowed gradually
from near the base to the acuminate, serrate apex, margins serrate, incised, pinnatifid
or pinnately divided, glabrous and striate above, under surface with some laciniate
scales near the base of the pinna and often on the veins ; veins numerous, crowded,
forked ; texture subcoriaceous. Sori linear-elongate, spreading along the veins
from the costa to near the margin. Spores plano-convex, 30-36 X 21, with a peri-

262 THE CEYLON SPECIES OF ASPLENIUM

spore forming a median undulate wing and irregular anastomosing thickened ridges
on the surface.

17a. Asplenium falcatum var. falcatum.

Fronds pinnate ; margins of pinnae serrate or shallowly to deeply incised with
toothed lobes.

CEYLON : Thwaites C.P. 1339 (BM ; E; GH; K; PDA). Gardner 1080 (CGE ;
K ; PDA—the Cambridge sheet localized as on rocks and trees in forests at Galle,
Apr. 1844). Common a few miles from Colombo and in many places in the hills,
Mrs. Chevalier (BM). Hulangamuwa, near Matale, Nov. 1861, Herb. Brodie (E).
Kurunegala Rock, 22 June 1927, Alston 703 (K ; PDA). Ascent and summit of
Ritigala, North Central Province, epiphytic, 22 Mar. 1905, Willis (PDA). Same
locality, 675 m., 14 Jan. 1951, Sledge 873 (BM; US). Between Ratnapura and
foot of Adam’s Peak, in forest at low elevation, 5 Jan. 1951, Manton in Sledge
838 (BM). Near Pahale Hewissa, Western Province, in forest, 75 m., 31 Mar. 1954,
Sledge 1375 (BM). Niriella, Sabaragamuwa Province, 225 m., 4 Feb. 1954, Schmid
1122 (BM)., 1899, Anderson (E). Ferguson (GH ; US 815487, 815488). Robinson
65(K). Walker 34(K). Wall(E; GL; K).

17b. Asplenium falcatum var. bipinnatum Sledge, var. nov.

Asplenium mysorense Roth, Nov. Pl. Sp. : 395 (1821) pro parte.

Tarachia furcata var. platyphylla C. Pres] in Abhandl. K. Bohm. Ges. Wiss., Folge 5, 6:
440 (1851) pro parte.

Asplenium praemorsum var. latum sensu T. Moore, Index Fil. : 156 (1859) pro parte ;
non Desv.

Asplenium spathulinum sensu Bedd., Ferns S. Ind.: 75, t. 226 (1864) ; non J. Smith
ex Hook.

Frondes bipinnatae ; pinnulae infimae pinnarum inferiorum et mediarum breviter
stipitatae vel subsessiles, rhomboideae, basi cuneatae, margine versus apicem
acutum serrata aliquando margine tenuiter vel profunde incisae, plerumque 1-5-3 cm.
longae, in exemplis magnis usque ad 6 cm. longae, cum apice attenuato ; pinnulae
mediae in basin posteriorem decurrentes, superiores connatae in apicem pinnae
serratum sensim mutatae ; pinnae supremae fere ut in var. falcato, rarius pinnatae
vel pinnatifidae.

CEYLON : Thwaites C.P. 2905 (E ; GH ; K ; PDA—the PDA sheets annotated :
Hantane, 1850 ; Haputelle, Apr. 1856, Apr. 1863). Thwaites C.P. 3891 (BM ; K ;
PDA—one of the PDA sheets annotated: Wattekelle, 1,200 m., June 1865).
Thwaites C.P. 1341 in part (PDA). Thwaites C.P. 3801 in part (PDA). Galle,
on trees and rocks in forests, Dec. 1844, Gardner 1083 bis (CGE). Summit of Riti-
gala, North Central Province, 24 Mar. 1905, Willis (PDA). Migahakihula, 270 m.,
Mar. 1890, Deschamps in Herb. Delessert (G). Nalanda near Matale, jungle by road-
side, 300 m., 13 Jan. 1951, Sledge 860 (BM, holotype). Pussalamankada, on the
Kandy-Maturata road, in forest, 540 m., 18 Jan. 1954, Sledge 983 (BM; US).
Weragamtota, by forest path, 150 m., 11 Feb. 1954, Sledge 1132 (BM). Near the
Maha Oya, south-east of Kandy, 450 m., 18 Feb. 1954, Sledge 1148 (BM). Beddome,
“type of Pl. ccxxv1 F.S.I.”’ (K). Mrs. Chevalier (BM). Ferguson (GH ; US

THE CEYLON SPECIES OF ASPLENIUM 263

815486, 815489). Gardner 1082 (PDA). Gardner 1084 (K). Macrae (E; W).
Wall (E; GL; K). Wight 1905 (E).
On rocks and trees at low or moderate elevations.

Madagascar, Mascarenes, Ceylon, India, south China, Malaya, Indonesia, Philip-
pines, Polynesia, Australia, New Zealand.

Linnaeus based his description of Tvichomanes adiantoides on a Ceylon specimen
collected by Hermann. The specimen is in the British Museum Herbarium (Herb.
Hermann, vol. 3, fol. 47). He also cited synonyms from Plukenet and Burman.
Lamarck based his Asplenium falcatum on specimens from Mauritius and included
the Burman synonym, and his is the correct name for this species since A. adiantoides
(L.) C. Chr., which has been widely employed for it, is an illegitimate combination as
Lamarck had used the name A. adiantoides for another species.

A. falcatum is widespread in tropical Asia and, inits typical form, is easily recognized.
It varies much in the size and form of the pinnae, the margins of which may be
subentire, serrate, irregularly incised, pinnatifid or pinnately divided. Clarke
(Trans. Linn. Soc. Lond., Ser. 2, Bot. 1: 480 (1880)) states that the fronds are
“sometimes pinnatifid nearly to the midrib” and Skottsberg (Medd. Géteb. Bot.
Tradg. 15 : 78-102 (1942)), who made a detailed study of variation in this and
allied species but who evidently saw little material from Ceylon, describes and illus-
trates the most extreme form as being no more than pinnatipartite. Other des-
criptions do not allow even this degree of lobing and none recognizes the existence
of bipinnate forms, though Compton (Journ. Linn. Soc. Lond., Bot. 45 : 446 (1922))
under A. adiantoides described a tripinnate variety from New Caledonia. Yet in Ceylon
fully bipinnate forms are not uncommon and a very instructive series of specimens
in the Kew Herbarium, collected by Wall, illustrates the wide range in form which
A. falcatum may assume from simply pinnate examples with serrate pinnae through
those with lobed and pinnatifid pinnae to fully bipinnate forms. Several of the
Peradeniya sheets have been labelled by Thwaites “Asl. falcatum forma composita ”’
and Baker has annotated one of Wall’s sheets at Kew “A. falcatum bipinnate
variety” though he never published a description. Some Ceylon plants transitional
between normal A. falcatwm and the var. bipinnatum agree with Skottsberg’s illus-
tration (tom. cit. : 79, fig. 211) of A. falcatum var. sectum (Hilleb.) Skottsb.

Although bipinnate forms of A. falcatum appear to be much more frequent in
Ceylon than elsewhere, they are not confined to Ceylon. Some specimens from
South India are identical with Ceylon plants of var. bipinnatum, and A. mysorense
Roth, described from Madras, consists at least in part of bipinnate fronds of A.
falcatum. Holttum (Fl. Malaya 2: 432 (1954)) refers to a Malayan specimen with
deeply dissected pinnae and a specimen at Kew from the same region (Bukit Ketri,
Perlis, 5 Feb. 1904, Matthew) is bipinnate. I have not seen the bipinnate variety
from any other region.

The middle and lower pinnae of fronds of A. falcatum var. bipinnatum look so
different from var. falcatum as to be scarcely recognizable as the same species. But
the distal end of the frond usually shows a transition to undivided pinnae of the

264 THE CEYLON SPECIES OF ASPLENIUM

normal type. Where, as is occasionally the case, the frond is bipinnate throughout,
all resemblance to the typical form is lost. Failure to recognize the relationship
between bipinnate fronds and the simply pinnate condition has led to confusion with
other species. The Ceylon specimen which Beddome described and illustrated
(Ferns S. Ind. : 75, t. 226 (1864)) as A. spathulinum from a specimen sent to him
under that name by Thwaites, is an example of A. falcatum var. bipinnatum. It
shows well the transition to the var. falcatum type of pinna in the distal part of the
frond. Ferguson (Ceyl. Ferns : 26-27 (1880)) was well aware from field experience
of the range of variation in the fronds of this species and he correctly identified
Beddome’s illustration as being bipinnate A. falcatwm, though he was incorrect in
also referring Beddome’s preceding illustration (tom. cit.: t. 225) to the same species.
Moore’s A. praemorsum var. latum includes bipinnate A. falcatum as well as A.
aethiopicum, and Macrae’s Ceylon specimen, included by Presl in his Tarachia furcata
var. platyphylla, is also A. falcatum var. bipinnatum.

Var. bipinnatum almost certainly covers a genetically heterogeneous group, for the
occurrence of large spherical spores or of abortive sporangia in some gatherings is
indicative of apogamous and hybrid strains, but such plants are not separable
morphologically from others having plano-convex spores similar to, though larger
than, those of var. falcatum.

The altitudinal range in Ceylon of A. falcatum is markedly different from that of
A. aethiopicum and A. affine, the two species with which the bipinnate variety has
most frequently been confounded. A. falcatum is a lowland fern whereas the other
two are ferns of high elevations. The highest altitude at which I have collected any
form of A. falcatum in Ceylon is 675 m., though Thwaites cites 4,000 ft. (1,200 m.)
on the label of a Kew specimen of the bipinnate variety, and the same altitude is
ascribed to an Indian specimen of A. falcatum var. bipinnatum also at Kew, collected
by Beddome at Coorg. A. aethiopicum and A. affine are both confined, in Ceylon,
to forests in the higher mountains of the interior. The former I have collected only
between 1,500 and 2,025 m. and the latter between 1,200 and 2,040 m.

18. Asplenium indicum Sledge, nom. nov.
Asplenium planicaule Wall., Numer. List: 8, n. 189 (1829), nom. nud.
Asplenium falcatum var. abbreviatum Kunze in Linnaea 24 : 260 (1851).
Asplenium planicaule Wall. ex Mett. in Abhandl. Senckenb. Naturforsch. Ges. 3: 201
(1859) ; non A. planicaule Lowe (1858).—Hook., Sp. Fil. 3 : 163, t. 200 B (1860).—
Bedd., Ferns S. Ind. : 47, t. 139 (1864).—Hieron. in Hedwigia 61 : 33 (1919).

Asplenium laciniatum sensu Bedd., Handb. Ferns Brit. Ind. : 154 (1883) pro parte ; non
D. Don.

Rhizome short, erect, apex clothed with dark-brown to black, narrow, tapering
scales. Stipes grey-green, up to 12 cm. long, scaly when young, as is the rhachis, with
linear-lanceolate, entire-margined, brown scales, becoming glabrous or almost so
with age. Lamina linear-oblong, up to 30 cm. long, 3-6 cm. broad, simply pinnate
with 12-20 pairs of pinnae, apex acuminate ; pinnae shortly petiolate, (I-) 2-4 cm.
long and nearly rt cm. broad in the widest part, dimidiate-ovate, acute, superior base

THE CEYLON SPECIES OF ASPLENIUM 265

broadly cuneate, subauriculate, inferior base narrowly cuneate and entire to 4 or more
the length from the base, the rest of the pinna above and below irregularly and shallowly
lobed with dentate margins ; veins close, once or twice forked ; texture stiff, sub-
coriaceous. Sori linear-elongate, spreading along the veins from near the costa but
falling short of the margin. Spores reniform to plano-convex, 39-45 x 24-30p,
with a narrow perispore wing and surfaces rugose with thickened anastomosing
ridges.

CEYLON: Thwaites C.P. 3289 (BM; PDA—data cited on PDA sheet: Pas Dun
Korle, Ramboda, etc., July 1847, Gardner; Hantane, Jan. 1854, 1863). Central Pro-
vince, I,200-1,500 m., Naylor Beckett 682 (BM). Vicarton, in forest, Wall in Herb.
Hance (BM). Nuwara Eliya, 1,500-1,800 m., Freeman 143 A(BM). Tangamalai
Sanctuary, Haputale, 1,500 m., 25 Feb. 1954, Sledge 1203 (BM). Kudaoya,
Ramboda Pass, among rocks by stream in jungle, 1,680 m., 16 Mar. 1954, Sledge
1294 (BM). Ferguson (GH ; US 815506). Walker (K). Wall (E). Wight (K).

On rocks and trees in forests above I,000 m. ; not common.

Réunion (teste Hieronymus), Ceylon, South India (Malabar mountains), Himalaya
from Kumaun to Assam (Khasi and Lushai Hills), China (Yunnan), Burma, Thai-
land, Indochina, Philippines, Japan (as var.).

Ceylon specimens match Wallich 18g, the type of A. planicaule Wall. ex Mett.,
which name, however, is a later homonym and has to be replaced. Hieronymus
(loc. cit.) has discussed this and the related species A. laciniatum, to which it had
been reduced as a variety by Christensen (Index Fil. : 126 (1905)), who did not, how-
ever, give it a varietal name.

A. indicum suggests a small form of A. falcatum with greatly shortened pinnae
and green stipes which are scaly when young. The texture and close venation of
the pinnae are similar to those of A. falcatum, to which it appears to be related
though, as Hooker (Sp. Fil. 3 : 163 (1860)) rightly said, no transitional specimens
occur which might be considered as a passage to that species. A. imdicum moreover
is a mountain species whereas A. falcatum is a fern of low altitudes, and the geo-
graphical ranges of the two species do not coincide.

19. Asplenium nitidum Swartz, Synops. Fil. : 84, 280 (1806).—Hook., Sp. Fil.

3: 172 (1860).—Tardieu-Blot & Ching in Notul. Syst. 5: 148, t. 1 fig. 1,
it. 2.lig. £ (1930);

Asplenium insigne Blume, Enum. Pl. Jav. 2: 188 (1828).

Asplenium spathulinum Kunze in Bot. Zeit. 6 : 524 (1848).

Tarachia nitida (Swartz) C. Presl in Abhandl. K. Bohm. Ges. Wiss., Folge 5, 6 : 443 (1851).

Asplenium glaucophyllum Alderw. van Rosenb. in Bull. Jard. Bot. Buitenz., Sér. 2, 7:

6 (1912).—Holtt., Fl. Malaya 2 : 440 (1954).

Rhizome short-creeping with clustered stipes, apex clothed with narrow, purple-
brown, wavy-edged, thin-walled, glossy scales up to 2 cm. long. Stipes stout, dark
coloured, glabrous; rhachis similar, grooved above. Lamina up to 100 X 30 cm.,
ovate to deltoid-ovate in outline, bipinnate, lowest pinnae not, or scarcely, reduced ;
largest pinnae to 15 X 8 cm. or more with up to 8 pairs of free pinnules below the

266 THE CEYLON SPECIES OF ASPLENIUM

lobed apex, pmnules variable in size and outline, sometimes short (about 2 cm. long)
and broad, rounded above and with serrate margins, or triangular and narrowed to
an acute apex, up to 7 cm. long and deeply lobed at the base with the broadly
rounded acroscopic lobe almost or quite free to the base, pinnules of upper pinnae
smaller with broadly cuneate to truncate upper base and narrowly cuneate lower
base, all striate and shining above, glabrous on both surfaces ; veins very numerous and
closely placed, in the larger pinnules springing from a fairly distinct costa ; texture
firm. Sori elongate, extending from near the midrib but falling short of the margin.
Spores “‘ with a wing of narrow to moderate width ”’.

CEYLON : ‘‘ Ceylon’”’, Mrs. Walker (K).
In lowland forests.

Ceylon, South India, Himalaya from Nepal to Assam, Malaya, Sumatra, Java,
Borneo.

The scales of A. mitidum are larger, lighter in colour and have thinner walls than
those of A. aethiopicum and A. falcatum : their edges are wavy and the surfaces
often more or less iridescent, as in the African A. friesiorum. The dark rhachis and
completely glabrous fronds, sometimes with a sheen on the upper surface, are
distinctive. It varies greatly in size. In the single Ceylon example at Kew the
largest pinnae are 14 x 8 cm., the largest pinnules 5 x 2 cm., narrowly triangular
in outline, pinnatifid at the base, dentate above and with acute apices. The specimen
matches closely Cuming 376 from Malacca, which Hooker correctly cited as typical
of the species ; this gathering is the type of A. spathulinum Kunze, who stated
erroneously that it came from the Philippines. Holttum uses the name A. glauco-
phyllum Alderw. van Rosenb. for this species, citing A. nitidum “ sensu Tardieu-
Blot et Ching”’ as a synonym, but as Tardieu-Blot & Ching’s description was
based on the type specimens in the Riksmuseum, Stockholm, which are illustrated
in their paper, their sense is surely also the original sense.

The specimens on which Swartz based his description of A. nitidum were collected
by Rottler at Tranquebar in Madras. The species might therefore be expected to
occur elsewhere in South India and Ceylon. But the specimen from Travancore
in Beddome’s collection at Kew, from which the drawing was made for Plate 149 of
his Ferns of Southern India, is the only gathering, other than Rottler’s specimens,
that I have seen from southern India, and the specimen and illustration, though
correctly referred to A. nitidum, do not represent a characteristic frond. There are
no specimens from South India or Ceylon in the British Museum collection. Mrs.
Walker’s specimen in the Kew Herbarium, collected a century and a quarter ago, is
the only example I have seen from Ceylon. There is no specimen in the Peradeniya
herbarium!. It is evident, therefore, that A. nitidum must be a very rare species
in both South India and Ceylon. It is also probable that it is still rarer now than

1 There is some doubt as to the provenance of the Kew specimen of A. nitidum despite the geographical
probability of its authenticity. Four other ferns in Hooker’s collection, labelled simply ‘‘ Ceylon ’’,
viz. Ctenopteris fuscata Kunze, Drynaria sparsisora (Desv.) T. Moore, Dryopteris khasiana C. Chr. and
Pyrrosia acrostichoides (Forst. f.) Ching, are similarly unrepresented in the herbarium at Peradeniya and

no other Ceylon gatherings are known. It may well be that all were sent to Hooker from plants grown
in the Peradeniya Botanic Garden.

THE CEYLON SPECIES OF ASPLENIUM 207

formerly, for it grows in forests at low elevations and these have been much reduced
in extent during the last century.

Thwaites and other Ceylon botanists were unacquainted with A. nitidum and
the specimens so named by them are A. affine, though Thwaites’s ‘‘ forma magis
composita. An A. laserpitifolium...C.P. 3801’ quoted under A. nitidum (Enum.
Pl. Zeyl. : 384 (1864)) isa large dissected form of A. aethiopicum. The fern Beddome
named A. laserpititfolium Lam. in his Ferns of Southern India (t. 225) and later, in
the Handbook, referred to A. nitidum, is also a form of A. aethiopicum.

20. Asplenium aethiopicum (Burm. f.) Becherer in Candollea 6: 23 (1935).
(Fig. 1.)
Trichomanes aethiopicum Burm. f., Fl. Ind., Fl. Cap. Prodr.: 32 (1768).
Asplenium adiantoides Lam. in Encycl. Méth., Bot. 2 : 309 (1786).
Asplenium praemorsum Swartz, Nov. Gen. & Sp. Pl.: 130 (1788).
Asplenium furcatum Thunb., Prodr. Pl. Cap.: 172 (1794).—Bedd., Handb. Ferns Brit.
Ind.: 157 (1883).
Tarachia furcata (Thunb.) C. Presl in Abhandl. K. Béhm. Ges. Wiss., Folge 5, 6 : 440

1851).
TeNae laserpitiifolium sensu Bedd., Ferns S. Ind.: 75, t. 225 (1864) ; non Lam.

Rhizome creeping, clothed with linear-lanceolate, dark-brown, glossy, hair-pointed
scales 5 mm. long. Fronds commonly 30-60 cm., sometimes 1 m. high. Stipes up
to 30 or more cm. long, purplish-black, usually more or less thickly clothed, as are
the rhachides and young fronds, with ferrugineous, hair-pointed, deciduous scales
intermixed with copious, slender, woolly, hair-like scales, but sometimes with few
scales more or less restricted to the base of the pinnae or practically glabrous.
Lamina bipinnate, 15-45 or more cm. long, 5-20 cm. broad, linear-lanceolate to
ovate-lanceolate with 12-25 pairs of shortly stalked, patent pinnae, the lowermost
somewhat reduced, the upper ones merging into the lobed apex ; pinnae 2-10 cm.
long, I-2:5 cm. wide near the base, acute or sometimes caudate, lower pinnules
rhomboid or subspathulate, base cuneate, apex truncate, rounded or acuminate,
split into two or three oblong, incised lobes with serrate apices; median and distal
pinnules broadly adnate, oblong with irregularly toothed apices ; glabrous or almost
so above, usually scaly or hairy beneath on the veins but sometimes glabrous ; .
veins close, flabellate, forked, giving a striate appearance to the upper surface of
the pinnules ; texture subcoriaceous. Sori linear, 0-5-1 cm. long. Spores plano-
convex, (39-) 41-45 (-48) x 24-27 (-30)u, with a perispore forming a median,
undulate wing and irregular, anastomosing, thickened ridges on the surface.

CEYLON : Thwaites C.P. 3497 (BM ; E ; K ; PDA—data on one of PDA sheets :
Badulla district, Feb. 1856; Horton Plains, Apr. 1856). Thwaites C.P. 3801 (BM;
K ; PDA). Thwattes C.P. 1341 (BM; K; PDA in part) (a short glabrous form
with broad flat pinnules). Hulangamuwa, Matale, Apr. 1858, Herb. Brodie (E)
(same form). Badulla, Freeman 162 E, 163 F (BM). Nuwara Eliya, Freeman 156 a,
157 B (BM). Same locality, 1830, Walker in Herb. Delessert (G). Moon Plains,
Nuwara Eliya, 1,800 m., 23 Dec. 1950, Sledge 7177 (BM; US). Pedrotalagalla,

268 THE CEYLON SPECIES OF ASPLENIUM

by path ascending mountain from Nuwara Eliya, 2,025 m., 26 Dec. 1950, Sledge
725 (BM). Hakgala, damp rocks on patana, 1,680 m., 5 Nov. 1897, Pearson 268
(CGE). Same locality, in jungle at 1,800 m., Dec. 1950, Manton (BM), Sledge 633
(BM). Same locality, 23 Dec. 1950, Holttum 39162 (SING). Sita Eliya, exposed
rock on patana, 1,740 m., 23 Oct. 1897, Pearson 875 (CGE). Between
Pattipola and Horton Plains, epiphytic in forest, 1,800 m., 20 Dec. 1950, Sledge
666 (BM). Above Ohiya, by path through forest to Horton Plains, 1,800 m., 30 Dec.
1950, Sledge 787 (BM). Le Vallon, in forest above tea estate, 1,500 m., 9 Feb.
1954, Sledge 1120 (BM ; US). Ramboda Pass, epiphytic in dense shade in forest
near the track to Maturata, 1,890 m., 17 Mar. 1954, Sledge 1302 (BM). Anderson (E).
Naylor Beckett 361 (GH). Beddome, “type of Pl. ccxxv F.S.1. Asplenium
laserpitiifolium ”’ (K). Ferguson (GH ; US 815490). Gardner 1082 (BM ; CGE;
K). Dec. 1876, Hutchison (E). Mrs. Walker (K). Wall (E).

In mountain forests above 1,500 m.
Tropical America, Africa from the Cape to Ethiopia and Madeira, Mascarene
Islands, Ceylon, South India, Java, Australia, New Caledonia, Hawaii.

This is a polymorphic species which varies much in size and degree of dissection of
the frond and in the degree to which the scaly clothing of the frond axis and pinnae
is developed. The fronds may be narrow in outline in tall as well as dwarf plants,
or they may be broad. In forms with narrow fronds the pinnae are either deeply
pinnatifid or pinnate ; wider-fronded plants have pinnate pinnae, the pinnules
being deeply pinnatifid or again pinnate. The pinnules may be closely or laxly
arranged on the pinna rhachis and may be narrow, the narrowness in dried specimens
often being accentuated by their revolute margins, or broad and flat.

The copious, ferrugineous, woolly, hair-like scales on the stipes and rhachides,
and often on the under sides of the pinnules, are normally characteristic and facilitate
identification. But forms occur in Ceylon, and elsewhere, in which these are absent
on mature fronds and such forms are more likely to be confused with other species.
Loss of scales is only partly due to age for fully mature fronds may remain woolly
whilst small immature fronds on other plants may be almost glabrous. Both in the
matter of indumentum and in frond form A. aethiopicum is undoubtedly a plastic
species. The silhouettes (Fig. 1) show fronds taken from the same rootstock in
successive years, the smaller frond being taken from the wild plant when collected,
the larger from the same rootstock after cultivation for one year at Kew. Apart
from the difference in size and dissection, the hair-like scales are almost confined
to the regions of the pinna bases in the larger frond. More extreme examples from
Ceylon of such large plants devoid of woolly indumentum have been referred to A.
laserpitufoium Lam. Beddome’s illustration (Ferns S. Ind.: t. 225 (1864), as
A. laserpitiifolium), drawn from a Ceylon specimen of C.P. 3801, shows such
an extreme form and it is hardly surprising that it should have been referred to another
species. The drawing shows an apparently glabrous plant, though there are in fact
some narrow scales with attenuated toothed apices on the specimen, which is pre-
served at Kew. The spore measurements of similar plants moreover agree with
those of A. aethiopicum. Some Nilgiri gatherings agree closely with such Ceylon

THE CEYLON SPECIES OF ASPLENIUM 269

plants in the form and cutting of the fronds, but the presence on them of many of
the characteristic hair-scales has facilitated correct identification, whilst their
absence on some Ceylon plants led Beddome, Thwaites and others to assign these
to another species.

Bipinnate forms of A. aethiopicum with broad, shallowly lobed pinnules and with-
out hair-like scales bear a close resemblance to bipinnate forms of A. falcatum.
Beddome’s illustration (Ferns S. Ind. : t. 144 (1864)) represents such a form and
should be compared with his illustration (tom. cit. : t. 226 (1864), as A. spathulinum)
which represents A. falcatum var. bipinnatum. The pinnae in the distal parts of the
frond, showing transitions in the one case to those typical of simply pinnate A.
falcatum, together with the larger spores and the more strongly creeping rhizome
with its narrow, hair-pointed scales in A. aethiopicum, afford means of separation.

The great variation displayed by A. aethiopicum is due partly to environmental and
partly to genetical factors. The specimen figured (Fig. 1) is known to be an octo-
ploid and other grades of polyploidy may well occur in Ceylon as tetraploid and
dodecaploid races are known from Africa. The large dissected form figured by
Beddome as A. laserpititfolium is similar to the African tetraploid. Some of my
Ceylon gatherings have abortive sporangia and may be of hybrid origin. A valid
subdivision can only be based on experimental studies, for any attempt to describe
a series of subordinate taxa to cover the wide range of variation in Ceylon would be
impracticable in view of the known fact that a wide morphological response to changes
in growth conditions or age may occur within an individual plant.

21. Asplenium affine Swartz in Schrad., Journ. Bot. 1800 (2) : 56 (1801).—

Sledge in Kew Bull. 15 : 401, fig. 2 (1962). (Fig. 2.)

Asplenium spathulinum J. Smith in Hook., Journ. Bot. 3 : 408 (1841), nom. nud.

Asplenium spathulinum J. Smith ex Hook., Sp. Fil. 3 : 170 (1860), nom. illegit., pro parte,

quoad specim. ex Luzon, Borneo et Ceylon; non A. spathulinum Kunze (1848).
Rhizome short-creeping, clothed with narrow, attenuate, dark-brown scales.

Stipes 10-20 cm. long (up to 40 cm. in forma majus), dark-coloured, sparsely scaly
when young, becoming glabrous or nearly so ; rhachis similar, grooved above.
Lamina bipinnate, 15-30 X 5-15 cm. (tripinnate at the base and up to 80 x 30 cm.
in forma majus), lanceolate to oblong-ovate in outline, often proliferous from buds
borne distally on the rhachis ; pinnae 10-20 on each side, lanceolate to ovate, acute,
the upper ones merging into the lobed apex ; largest pinnae up to 8 x 3 cm., bearing
I—5 pairs of spathulate or broadly ovate pinnules below the lobed apex ; finnules
usually blunt and rounded with toothed apices—more rarely subacute—and often
auriculate on the anterior margin, the largest 1-2 cm. long, entire or shallowly lobed
or sometimes with the anterior auricle free almost to the base ; surfaces glabrous
or with a few narrow scales on the rhachides beneath ; veins forked, not crowded,
basal acroscopic pinnules showing 5-18 ultimate vein endings according to size ;
texture chartaceous. Sori elongate, falling short of the margin. Spores reniform
to plano-convex, 36-57 24-30u, with a conspicuous undulate perispore wing and
anastomosing surface folds.

270 THE CEYLON SPECIES OF ASPLENIUM

21 (i). Asplenium affine forma affine.

Fronds of moderate size, 30-60 X 10-15 cm., lanceolate ; pinnae up to 8 cm. long,
bearing I-5 pairs of spathulate or broadly ovate pinnules with rounded, toothed
apices and often auriculate on the anterior margin ; largest pinnules 1-2 cm. long,
entire or shallowly lobed or sometimes with the the anterior auricles free almost
to the base.

CEYLON : Thwaites C.P. 1008 (BM in part ; E ; GH.; K ; PDA in part—data
on PDA sheet : Ramboda, Hantane, etc., 1847, Gardner). Thwaites C.P. 2905 in
Herb. Hance (BM). Nuwara Eliya, Hooker & Thomson 176 (K). Same locality,
on banks and in woods, 1,800 m., 9 May 1906, Matthew (K). Same locality,
Freeman 159 B, 161 D, 164 G, 165H (BM). Same locality, 1,950 m., 24-27 Feb. 1954,
Schmid 1359 (BM). Horton Plains, in jungle near the rest house, 2,040 m., 19 Dec.
1950, Sledge 699 (BM ; US). Same locality, epiphytic in mossy forest, c. 2,100 m.,
30 Dec. 1950, Holttum 39223 (SING). Hakgala, in jungle, 1,800 m., 16 Dec. 1950,
Sledge 639 (BM). Same locality, 23 Dec. 1950, Holttum 39172 (SING). Tangamalai
Sanctuary, near Haputale, 1,500 m., 25 Feb. 1954, Sledge 1216 (BM ; US).
Namunukula, Freeman 158 A (BM). Tonacgmbe Estate, Namunukula, terrestrial
in jungle, 1,350 m., 23 Feb. 1954, Sledge 1175 (BM). Le Vallon, in jungle above tea
estate, 1,500 m., 9 Feb. 1954, Sledge 11177 (BM). Oodawella, near Kandy, in jungle
above tea estate, 1,200 m., 8 Dec. 1950, Sledge 532 (BM). Corbet’s Gap, in jungle,
1,200 m., 7 Jan. 1951, Sledge 850 (BM). Matale, Fraser (CGE). Badulla, Herb.
Brodie (E). Naylor Beckett 27 (E ; GH). Mrs. Carr in Herb. T. Moore (K). Mrs.
Chevalier—on sheet with two specimens of A. falcatum var. bipinnatum (BM).
Ferguson (GH ; PDA). Gardner 1083, 1084 (E). Gardner in Herb. J. Smith (BM).
1876, Hutchison (E). Walker 106 in Herb. Delessert (G). Walker (K). Wall (E ;
GHeiGL).

21 (ii). Asplenium affine forma majus Sledge in Kew Bull. 15 : 408, fig. 3 (1962).

Asplenium affine sensu Hook., Sp. Fil. 3 : 169, t. 202 (1860) ; non Swartz.

Asplenium nitidum sensu Bedd., Ferns S. Ind.: 50, t. 148 (1864) ; non Swartz.

Asplenium cuneatum var. affine Baker, Fl. Maurit. & Seych.: 489 (1877) pro parte. ;

non A. affine Swartz.
Fronds large, up to 1 m. long and 30 cm. wide, broadly lanceolate ; pinnae more

attenuate or even caudate, up to 18 cm. long with 8-9 pairs of ovate-acute pinnatifid
or pinnate pinnules with toothed margins, the largest 3-4 (—5) cm. long.

CEYLON : Thwaites C.P. 1008 (BM in part ; PDA in part). Hantane range,
on trees in forests, Oct. 1844, Gardner 1083 (CGE). Galleheria, Thwaites in Herb.
Beddome (K). Nuwara Eliya, Freeman 160 c (BM). Hakgala, jungle below road,
22 May tro11, J. M. Silva (PDA). Same locality, terrestrial in forest, 1,650 m.,
26 Feb. 1954, Sledge r210 (BM). Ex hort. Kew., 1956, Alston 77634 (BM). Ferguson
(GH ; US 815504).

21 (ili). Asplenium affine forma tenuisectum Sledge, forma nov.

Pinnulae angustae circa 2 mm. latae, eae basales acroscopicae profunde pinnatifidae
vel pinnatae cum segmentis angustis.

THE CEYLON SPECIES OF ASPLENIUM 271

CEYLON : Thwaites C.P. 3801 in part (PDA, holotype).
Common in forests above 1,200 m.

Madagascar, Mascarene Islands, Seychelles, Ceylon, South India, Hainan, Thailand,
Cambodia, Malaya, Sumatra, Borneo, Philippines, New Guinea, New Hebrides, Fiji.

A. affine was described from a Mauritius specimen which I consider inseparable
from Ceylon plants. Plants from the Seychelles so identified by Christensen are
certainly identical with Ceylon specimens. A similar specimen, Cuming 210 from
the Philippines, was named A. spathulinum by John Smith but was not described.
Hooker validated this name by description and A. spathulinum J. Smith ex Hook.
has been widely used for supposedly distinct plants from south-east Asia. In my
paper on A. affine and A. spathulinum (Kew Bull. 15 : 401-410 (1962)) I cited
Cuming 210 as the holotype of the latter name. But as Mr. C. V. Morton has pointed
out (im litt.) Hooker’s description was drawn from several collections including the
type collection of A. imsitictum Brackenr. which is cited as a synonym. As this had
already been validly described, A. spathulinum J. Smith ex Hook. is synonymous
with, and a superfluous renaming of, the Hawaiian A. insiticium Brackenr. More-
over, A. spathulinum J. Smith ex Hook. is in any case a later homonym of A. spath-
ulinum Kunze (Bot. Zeit. 6: 524 (1848)) and is therefore doubly illegitimate.
Kunze’s species was based on Cuming 376 from Malacca (not the Philippines as stated
by Kunze and Mettenius). Mettenius (Abhandl. Senckenb. Naturforsch. Ges. 3: 204
(1859)), Hooker (Sp. Fil. 3 : 173 (1860)) and Tardieu-Blot & Ching (Notul. Syst.
5: 148 (1936)) all refer Cuming 376 to A. nitidum Swartz and the sheet of this
number at the British Museum bears out their identification.

In my paper referred to above I have shown that A. affine Swartz and A. spath-
ulinum auct. cannot be maintained as separate species. A. affine is a very variable
species and fronds taken from the same rhizome before and after cultivation (see
Fig. 2) display considerably greater differences than those by which it has been
customary to distinguish it from A. spathulinum auct. Wild specimens are frequently
met with similar to the large-fronded form which I have distinguished as A. affine
forma majus, and these have often been assigned to different species. It was on
such a robust form that Hooker based his description and figures of A. affine, con-
sidering it—not unnaturally—to be specifically distinct from Smith’s Philippine
A. spathulinum. So arose the belief that two species exist, one in the Mascarene
Islands and one in the south-eastern Asia and the islands beyond ; a belief which
has been upheld by recent writers, though specimens available in herbaria give no
support to this view. A. affine forma tenuisectum also merits recognition as an
extreme variant, for its narrow pinnules are so different in appearance from those of
typical A. affine that the relationship is not obvious. Octoploid and dodecaploid
races of A. affine exist in Ceylon but apart from the greater average spore size in
the latter they do not appear to be characterized by any constant morphological
differences which will serve to separate two taxa.

Thwaites and other Ceylon botanists were unacquainted with the true A. nitidum
and used this name for plants of A. affine. Specimens of A. affine were distributed
by Thwaites, C.P. 1008, as A. nitidum. His A. spathulinum was partly A. falcatum

272 THE CEYLON SPECIES OF ASPLENIUM

var. bipinnatum and partly a form of A. aethiopicum. <A specimen of A. falcatum
var. bipinnatum sent by Thwaites to Beddome as A. spathulinum is figured over the
latter name on plate 226 of his Ferns of Southern India. Later, in his Handbook, Bed-
dome referred this illustration to A. affine, adding the comment “probably taken
from a specimen of furcatum and not from affine as represented at Kew’’. He thus
gave it three different names and erroneously in each case. Ferguson also referred
specimens of A. affine to A. nitidum and used the name A. affine for specimens of
A. falcatum var. bipinnatum. The fern figured by Beddome on plate 148 of his
Ferns of Southern India as A. nitidum represents A. affine forma majus and should be
compared with those figured by Hooker (Sp. Fil. 3 : t. 202 (1860)) and Sledge (Kew
Bull. 15: 407, fig. 3 (1962)). There is no representation of typical A. affine in
Beddome’s illustrations but silhouettes of fronds from six different plants are shown
in my paper (tom. cit. : 404, fig. 2) together with a silhouette (tom. cit.: 402,
fig. 1) of the type specimen.

Despite the confusion which has surrounded the use of the names A. affine and A.
nitidum in Ceylon, the former is a well-marked species quite distinct from members
of the A. falcatum-A. aethiopicum-A. nitidum group. In A. affine the fronds are
often proliferous from a bud arising on the distal part of the rhachis. Normally
the bluntly rounded, broadly ovate to spathulate pinnules of A. affine are distinctive,
although all four species are subject to considerable variation as regards degree of
dissection of the frond. The veins in the pinnules of A. affine, however, are always
relatively few in number and fairly widely spaced, basal acroscopic pinnules usually
showing from 5 to 18 ultimate vein endings according to size. In A. falcatum var.
bipinnatum, A. aethiopicum and A. nitidum the veins are repeatedly forked and much
more closely spaced, comparable figures for basal acroscopic pinnules being 20-50
and sometimes in large specimens of A. nitidum exceeding 100. A. affine is also a
fern of mountain forests whereas A. nitidum, of which I have seen only a single
specimen from Ceylon, and A. falcatum var. bipinnatum are both lowland plants.

22. Asplenium varians Hook. & Grev., Ic. Fil. 2: t. 172 (1829).—Hook., Sp.
Fil. 3 : 192 (1860).—Bedd., Ferns S. Ind. : 44, t. 129 (1864) ; Handb. Ferns
Brit. Ind.: 158 (1883).—Hope in Journ. Bombay Nat. Hist. Soc. 13: 667, t. 20
(1901).

Rhizome erect, clothed at the apex with linear, hair-pointed, dark-chestnut
scales. Fronds tufted, up to 20 cm. long and 1-5—4:5 cm. broad, oblong-lanceolate,
the lower pinnae not much reduced. Stipes slender, green, with filiform scales
when young, glabrescent. Pinnae 8-12 on each side, r—3cm. long, pinnate or
deeply pinnatifid, obtuse ; pinnules few, rounded with acutely toothed outer edges
and cuneate bases, glabrous above and beneath like the slender green rhachis ;
texture thin, herbaceous ; veins forked. Sori 2-4 on each pinnule ; indusium pale,
membranous, entire. Spores plano-convex, 27-30 X 2I-24yu, with a perispore
forming a reticulate network of surface folds.

CEYLON : Thwaites C.P. 3139 (BM; E; GH; K ; PDA—data cited on PDA
sheet : Uva, Apr. 1854). Hakgala, 1,500 m., Aug. 1881 (PDA). Ramboda, Wall in

THE CEYLON SPECIES OF ASPLENIUM 273

Herb. Hance (BM). Nuwara Eliya, Freeman 166 A, 167 B, 168 c, 169 D (BM).
Badulla road, below Hakgala, 1871, Robinson (K). Namunukula, in stony
ground and in crevices of wall by path through tea estate, 1,575 m., 24 Feb.
1954, Sledge 1199 (BM ; K ; US). Ferguson (GH). Wall (E ; GH).

In crevices of walls and under rocks in the elevated parts of the interior, goo-
1,600 m.

South Africa, Ceylon, India, China, Japan, Hawaii.
This is a small fern superficially resembling Athyrium hohenackeranum more than
any other Ceylon species of AsPlenium.

23. Asplenium disjunctum Sledge, sp. nov. (Fig. 3.)

Rhizoma erectum, apice paleis lineari-lanceolatis acutis integris atrofuscis c. 3 mm.
longis vestitum. Frondes fasciculatae, usque ad 20 cm. longae, 5 cm. latae. Sti-
pites 4-8 cm. longi, griseo-virides vel brunnescentes, juventute, ut rhachides,
squamulis patentibus angustis acutis brunneis instructi, denique denudati ; rhachis
sursum sulcata, im parte superiore sed infra apicem gemmam nigro-squamulosam
proliferantem gerens. Lamina ambitu anguste oblongo-lanceolata, bipinnata,
iripinnatifida, segmentis angustatis 1-2 mm. latis, pinnis circa 10-jugis, infra apicem
simpliciter pinnata, infimis vix brevioribus, marginibus pinnarum saepe in sicco
revolutis; pinnae breviter petiolatae, 1 cm. inter se remotae, ad 3 X 1-5 cm., ambitu
ovatae ; pinnulae 2~4 utroque latere, infra apicem pinnatifidae, basales pinnarum
inferiorum et mediarum profunde pinnatifidae, segmentis angustis integrisque, supra
glabrae, infra squamulis paucis fusco-cellulis basi lacerato-fimbriatis apice longe
attenuatis praesertim juxta basin pinnarum conspersae ; venae in segmentis semel
vel bis furcatae, in sicco supra manifestae ; textura firmiter herbacea. Sori in
extrema parte venarum ad apicem segmentorum attingentes. Sporae plano-convexae,
33-36 X 24u, perisporio juga spissata et anastomosantia formanti.

CEYLON : Passara, Uva Province, 450 m., Feb. 1885, Pole (K ; PDA, holotype).
Mandagala, Eastern Province, May 1885, Nevill (PDA). Ferguson (PDA).

Eastern side of the island at low elevations.

Endemic.
This is the fern referred to by Beddome (Suppl. Ferns Brit. Ind. : 31 (18g2)) as
probably an abnormal variety of “ Asplenium furcatum”’. He probably saw only

the single very indifferent specimen which is preserved in his collection at Kew.
There is in my opinion no relationship with that species or with any other Ceylon
species. Its finely cut, proliferous fronds give it some superficial resemblance to
A. tenuifolium ; but in A. tenuifolium the larger tripinnate-quadripinnatifid, glab-
rous fronds are membranous in texture, light-green in colour with green stipes and
rhachides, and the ultimate segments are sharp-pointed with solitary immersed veins
which terminate well below the tips of the segments.

A. disjunctum seems to me to be most closely allied to A. elmeri Christ, which was
described from the Philippines and has since been found in Borneo. From this it

| mm.

BIGs3:

THE CEYLON SPECIES OF ASPLENIUM

~S> \y//
404 Ns

Ne

Asplenium disjunctum Sledge (holotype). a, whole plant; 0, frond tip with gemma;
c, underside of pinnule ; d, rhizome scale ; é, spore.

THE CEYLON SPECIES OF ASPLENIUM 275

differs in facies and in the position of the sori, which are always situated at the ends
of the segments. The pinnules are narrower, less markedly cuneate and more
sharply incised, the narrowness of the pinnules and their segments being accentuated
by the tendency for their margins to be revolute in the dried specimens, a tendency
which is not apparent in specimens of A. elmert. The latter species is known only
from wet forests on high mountains in Luzon (Mt. Benguet) and Borneo (Mt. Kinabalu)
at altitudes of 1,500-4,000 m. Both the stations for the Ceylon species are at low
elevations on the eastern, dry side of the island.

The name refers to the separate and distant stations, 50 miles (80 km.) apart,
whence the two localized gatherings weremade. That from Mandagalain the Eastern
Province is far removed from any other station whence I have seen collections.

24. Asplenium tenuifolium D. Don, Prodr. Fl. Nepal. : 8 (1825).—Bedd.,
Ferns S. Ind. : 44, t. 130 (1864) ; Handb. Ferns Brit. Ind. : 159 (1883).

Rhizome oblique, paleaceous with narrow, finely pointed, dark-brown scales.
Fronds tufted, tripinnate or quadripinnate, normally 30-40 x 6-12 cm., sometimes
larger, lanceolate or oblong-lanceolate, the lowermost pinnae slightly reduced.
Stipe and rhachis slender, grooved above, glabrous or with a few hairs. Lamina
green, 10-30 cm. long, with 10-15 pairs of alternate, stalked, patent pinnae, the
lower ones 5~7 cm. long, lanceolate, bipinnate to tripinnate, proliferous from scaly
buds in the upper part, ultimate segments narrowed below and with acute to mucronate
tips, apices of the frond and primary pinnae pinnatifid with linear segments ;
glabrous above and beneath ;_ veins solitary, central, terminating well below the
apices ; texture thinly herbaceous. Sori 1-2 to a segment; indusium pale, mem-
branous, entire. Spores reniform to plano-convex, 30-36 xX 21I-27u, with a peri-
spore forming a broad undulate wing and many anastomosing surface folds.

CEYLON : Thwaites C.P. 3268 (BM; E; GH; K ; PDA—data cited on PDA
sheet : Nuwara Eliya, Apr. 1856). Hakgala, Aug. 1881 (PDA). Nuwara Eliya,
Robinson (K). Same locality, Ferguson (PDA). Same locality, Freeman 172 Cc
(BM). Same locality, 1,950 m., rr Mar. 1954, Schmid 1492 (BM). Blackpool,
near Nuwara Eliya, Freeman 170 a, 17% B (BM). Ramboda Pass, on rocks by
stream in forest on the Maturata track, I,920 m., 17 Mar. 1954, Sledge 1306 (BM;
K; US). 1899, Anderson (E). Ferguson (GH; US 815517). Gardner 1079
(K). Wall (E; GH).

On rocks by streams about Nuwara Eliya.

Ceylon, South India, North India from Nepal to Assam, west China (Yunnan),
Thailand, Indochina, Formosa.

This is a very distinct species on account of its finely dissected fronds. Hooker,
Beddome and Clarke do not refer to the proliferous habit though many Indian
specimens show small scaly buds, often dormant. In Ceylon specimens the fronds
invariably bear buds, often two or three on each frond, which arise in the groove
at or close to the junction of the pinna rhachis and first acroscopic pinnule.

276 THE CEYLON SPECIES OF ASPLENIUM

25. Asplenium decorum Kunze in Bot. Zeit. 6 : 176 (1848).

Darea belangeri Bory in Bélanger, Voy. Ind.-or., Bot. 2 : 51 (1833).

Asplenium belangeri (Bory) Kunze, loc. cit. (1848) ; non A. belangeri Bory (1833).—
Bedd., Handb. Ferns Brit. Ind.: 161 (1883).—Hboltt., Fl. Malaya 2 : 434 (1954).

Asplenium veitchianum T. Moore, Index Fil.: xlix (1857), 176 (1860).

Asplenium prolongatum Hook., Sp. Fil. 3 : 209 (1860) pro parte, quoad specim. Zeyl.—
Thw., Enum. Pl. Zeyl.: 384 (1864).

Rhizome erect, scaly at the apex, scales ovate-acute, black with paler, brown
edges. Fronds tufted, up to 30 cm. long. Stipes green, up to 10 cm. long, scaly
below with narrow, dark, clathrate scales which are often jagged at the base, inter-
mixed with and replaced higher on the stipe and on the under side of the rhachis and
pinnae by scattered, pale-brown, appressed, irregularly laciniate to stellate-
squamulose scales. Lamina pale-green, up to 20 cm. long, rarely more, 4-6 cm.
broad, oblong-lanceolate to ovate in outline, apex acute or sometimes caudate,
bipinnate ; midrib of rhachis raised above and bordered by narrow lateral wings ;
pinnae 2-5 cm. long, linear-oblong cut down regularly into numerous (up to 12 pairs)
linear, straight or somewhat falcate segments 1 mm. broad, the posterior margins
decurrent to form a narrow wing to the pinna rhachis, the lowest acroscopic segment
bifurcate, the rest simple and traversed by a single central vein; texture somewhat
fleshy. Sori one to each segment on the anterior margin ; indusium whitish, not
reflexed at maturity. Spores light-brown, plano-convex, 39-42 X 24-27u, with a
perispore wing and surface folds.

CEYLON : Thwaites C.P. 1348 (BM; E; GH; K; PDA). Kanneliya Forest
Reserve, Southern Province, 150 m., 2 Apr. 1954, Sledge 1386 (BM). Beddome (BM).
Ferguson (GH ;_ US 815511, 815512). Lowcountry forests, Hutchison (E). Wall
(E + GL). 1837; Wight 132 (E).

Epiphytic in wet forest in the south, especially (? exclusively) the Sinha Raja forest.

Ceylon, Indochina, Malaya, Sumatra, Java, Borneo, Celebes, New Guinea, Philip-
pines.

Ceylon plants were referred by Hooker to A. prolongatum Hook. and were so
named by Thwaites. They differ however in several respects. They lack the
remarkably attenuated apical prolongation of the rhachis, they never bear a terminal
bud, the shape of the frond is oblong-lanceolate to ovate, not linear-oblong, the
pinnae are sparingly furfuraceous or stellate-squamulose beneath and their divisions
are shorter but more numerous, and the basal scales are not so long or attenuated.
Beddome’s illustration (Handb. Ferns Brit. Ind.: 162, fig. 79 (1883), as A. rutifolium)
which represents typical A. prolongatum well illustrates the difference between this
species and A. decorum. Sometimes the apex of the frond is drawn out or caudate,
but it then bears simple, oblique and distant lobes and lacks the sudden transition
from pinnae to long naked extensions bearing terminal bud swhich are so character-
istic of A. prolongatum. The qualification in Hooker’s statement (Sec. Cent. Ferns:
sub t. 42 (1861)) that “‘ Nearly every one exhibits the remarkable prolongation of
the rachis, rooting at the apex ’”’ was evidently inserted to cover the few Ceylon
specimens which were seen by him, as all the other gatherings he quoted (and

DHE CEYLON SPECIES OF ASPLENIUM 277

many subsequent gatherings correctly referred to this species) are quite consistent in
this character. Christensen evidently doubted that the Ceylon plant was identical
with A. prolongatum as he excluded Ceylon from the distribution of that species as
given in the third supplement to the Index Filicum.

The African A. rutifolium (Berg.) Kunze, with which Ceylon plants were at one
time identified, differs in its ovate or lanceolate pinnae, the basal acroscopic pinnules
of which, and often the basiscopic ones also, are enlarged and pinnate.

The epithet belangert which has been widely used for this species is not available as
A. belangeri (Bory) Kunze is a later homonym of A. belangert Bory (= A. longissimum
Blume). Moore pointed this out long ago (Index Fil: xlix (1857), 176 (1860)) and
substituted the epithet vertchianum. But Kunze had already described A. decorum,
based on a plant from Java (Zollinger 1260), which he considered specifically distinct
from his A. belangert. The differences on which it was founded however are trivial
and all recent workers have agreed in recognizing only one dareoid species in Malaysia,
under which A. decorum has been cited as a synonym.

This species requires further study. Whilst I am sure that the Ceylon population
cannot be included under A. prolongatum Hook., I am by no means sure that it is
identical with Malaysian A. decorum. Plants from Malaya and Indonesia are mostly
considerably larger than those from Ceylon and have more regularly divided pinnae ;
but the frond apex is the same in both and although Malaysian plants frequently
produce buds they are not invariably proliferous as in A. prolongatum.

A. decorum has often been regarded as a variety or form of A. ¢enerum differing
only in its deeply and regularly pinnatifid pinnae. Although Holttum regards it as
a distinct species, he considers it closely related to A. tenerum, with which it frequently
grows in Malaya. In Ceylon, however, it is geographically separated from A.
tenerum, which is confined to the mountainous region of the interior, A. decorum
occurring only in the south of the island, and at lower elevations.

Bull. B.M. (N.H.) Bot. 3, 6 PLATE 20

Srorsm anret dantbich efach obey
oye bn ton Did he aan Pmwvnee + tome
‘ tee weg eee ne

Aeesete: wed edency wana. See hanheceton. :

Asplenium erectum Willd. (Bory 70, holotype.)

INDEX TO VOLUME*3

The page numbers of the principal references and the new taxonomic names are printed in bold-face

type, synonyms in italics.

Names of infraspecific taxa are indexed only when they are new or are

synonyms.
Acalypha annobonae 113 Anthocodium fragile ; ; : - 196
ACANTHACEAE 108 APOCYNACEAE : ; : : aL LO!
Achnanthes andicola 85 ARACEAE ‘ ; : ; : s- 116
baldjickii 81 Arachis hypogaea . : : : eos
subsp. lorenziana 82 Aralia guilfoyler. : F : » 102
var. subquadrata 82 ARALIACEAE ; : ; : 5 LOZ
baldjickii 82 Artocarpus communis. : , . 113
danica 76 heterophyllus. : s : . 113
var. maxima 72 integer 3 ‘ : : > E13
danica 2 Oe Arum sagittifolium ; : : . 116
fimbriata 72 ASCLEPIADACEAE . : : : 1 LO4
heteropsis 72 Asclepias curassavica. : 104
hungarica . 84 Asplenium, The Ceylon Species: of 233- 277
lorenziana . oi igtee Asplenium . : ; : : . 240
lyvata we 79 adiantoides : : : . 20%,.267
manifera 72 aethiopicum : ‘ ; : . 267
reichardtiana 70 affine : : : . 269
stvoemii G2 forma tenuisectum. ; : . 270
vaszaryi F 72 affine : : ; : : e270
Achnanthidium baldjickii 81 belangeri . ‘ : 3 : ) 270
hungaricum ‘ 84 brachyotus 3 : : : 252
neglectum 84 brasiliense : : : Z . 248
Achyranthes aspera 109 camptorhachis . : : : «248
Actinoneis lorenziana 82 caudatum . : : : . 258,259
Adenopus breviflorus 101 cheilosorum ; ‘ c : . 245
Adenostemma perrottetii 103 contiguum F ; : : . 259
Agelaea ovalis 98 coriaceum . : : ; , 255
Aghardia adhaerens 197 cristatum . : : : + 247
Alectra communis 107 cuneatum var. affine : : . ae],
sessiliflora 107 decorum . ; : : : . 276
Allamanda cathartica 104 decrescens . : : : ; . 259
Alternanthera maritima . 109 disjunctum : : : : . 273
sessilis 109 elongatum . : ; : ‘ 253
AMARANTHACEAE 109 emarginatodentatum . : eas
Amaranthus spinosus 109 ensiforme . : ; : : . 242
viridis : 109 erectum . : : ; ; . 248
AMARYLLIDACEAE . 115 evosodentatum ; 3 ‘ Sa eAT
ANACARDIACEAE 98 erythrocaulon. , : ; 245
Andropogon brevifolius : 118 excisum 3 : F . . 2406
Angiosperms of the Cambridge Annobon falcatum . : ; ; . 261
Island Expedition 93-118 var. abbreviatuis : : : 3 204
Angraecum talbotii . 115 var. bipinnatum . : : . 262
Annobon Island Expedition, Angiosperms formosum . : : ; : . 243
of the Cambridge 93-118 furcatum . fi ; F ‘ = 267
Annona muricata > 96 gardneri. ‘ : é ; . 258
reticulata . 96 glaucophyllum . : é : . 265
ANNONACEAE 96 heterocarpum ‘ ; : ; . 2245
Anthephora cristata 117 hirtum : ‘ : ; a. 254

280

Asplenium—conid.
inaequilaterale
indicum
Insigne
laciniatum .
laetum
laserpititfolium
longipes
lunulatum .
var. camptorhachis
var. evectum
var. sphenolobium
var. tvapeziforme
macraei forma sphenolobium
macrophyllum
minus
multijugum
mysovense .
nidus
nitidum
nitidum
normale
obscurum .
opacum
pavonicum .
pellucidum
persicifolium var. laoham :
planicaule .
praemorsum
var. latum
prolongatum
pyvamidatum
vesectum
serricula
servviforme .
spathulinum
sphenolobium
tenerum
var. terminans
teneyum
tenuifolium
trapeziforme
unilaterale
forma majus
var. majus
varians
vettchiorum
vulcanicum
walkerae
wightianum
zenkeranum
Avicennia africana
germinans .

Ball, P. W., and V. H. Heywood
Bambusa vulgaris . :
Bangia fuscopurpurea

BANGIACEAE .
Basella alba .

INDEX

BASELLACEAE : : . 2 2 EO
252 Beckeropsis Jaxior : ; ‘ . S18
264 nubica : : : : > “ris
265 Bertiera annobonensis : : ‘ . 102
264 Blidingia minima : : . 180,183
245 Boerhavia coccinea ; : ‘ . 108
267 diffusa ; : : , : - 109
256 diffusa : : : : ‘ . 108
248 Bolusiella talbotii . . 2 ; eet Ey
248 Bostrychia mixta ; : é 179,227
248 Bryonia palmata . : 2 : ~/ SIO
248 scabra : ‘ : : PLO?
252 Bryonopsis laciniosa ; IOL

248 Buchenavia and Ramatuella, N revision of
258 the genera . F , ; ; . I-46
243, 248 Buchenavia . : ‘ é ; : 4
243 acuminata 2 : : ; » «ao
262 callistachya : ; ; : ~ -20
242 capitata . : : : F see?
265 congesta . : ; ; : . 36
270 corrugata . : : : : : 19
243 discolor. : : ; F ee
247 fanshawei . 4 : : . eee
243 fluminensis : ; : : ese
243 gracilis : : ‘ ; : SY)
254 grandis. : ; ; : » oo
254 huberi : : ; ; : . 34
264 kleinii ; : , ‘ = : 14
267 longibracteata . : : : . 33
262 macahensis ; : ‘ : : 37
276 macrophylla F é : : ey
248 megalophylla. : ; : . 26
245 ochroprumna é : . : : 16
255 oxycarpa . : ‘ : : . 24
- 247 oxycarpa . : : : : - 2I
262, 205, 269 pallidovirens : : 5 : Sey 7/
248 parvifolia . 2 : : : o&S
253 ptariensis . ‘ : : 3 a ee
253 pterocarpa : : ‘ : ee
258 pulcherrima . : : : oe
275 punctata . ; 3 ‘ : ae 3
252 reticulata . ; 3 j : ‘ 17
245 sericocarpa ; ‘ , : . 25
246 stellae c : ; . é <2 132
246 suaveolens , : : : . 21
272 tomentosa . : : Fi j 2 ld
276 vaupesana : : : : oe EZ
256 viridiflora . : ; : : . 28
256 Bucholzia maritima : : : . 109
255 Bucida angustifolia , : ‘ 3, a pk
254 capitata. : ‘ : oar iy
108 Buforrestia imperforata ‘ : : ee cde
108 Bulbophyllum melanorrhachis : - 215
Caesalpinia pulcherrima . : : a
119-172 Cajanus cajan : ; ‘ ‘ . 9
penile) Calonyction aculeatum ; A . 105
179, 208 Calothrix crustacea 179, 182
208 CAMPANULACEAE . P : : 4.” JEG
110 Canscora decussata ‘ ; ‘ . 105

INDEX

Capartogramma, The diatom genus, and

the identity of Schizostauron
Capartogramma
amphoroides
crucicula .
jeanit
karstenii .
rhombicum
CAPPARACEAE
Capsicum annuum .
Carica papaya
CARICACEAE .
Caryophyllus avidus
Cassia obtusifolia
occidentalis
tova
Cassipourea annobouenss
Catharanthus roseus
CELASTRACEAE :
Centroceras clavulatum .
Cephalostigma perrottetii
CERAMIACEAE :
Ceramium clavulatum
Ceylon species of Asplenium, The
CHAMAESIPHONACEAE
Chamberlain, Yvonne M.
CHLOROPHYTA
Chorda lomentaria .
Citrus aurantifolia .
Cladophora catenifera
radiosa
‘ rupestris
CLADOPHORACBAE .
Clayton, W. D.
Cleistanthus libericus
Cleome ciliata
rutidosperma
Chitoria alba .
Coccochloris deusta .
Cocconeis beltmeyeri
danica
delicata :
diaphana var. Bp
dirupta
flexella
hungarica .
CoDIACEAE
Codiaeum variegatum
Codium adhaerens .
fragile é
Coffea liberica
Coix lacryma-jobi .
Colpomenia sinuosa
COMBRETACBAB :
Commelina benghalensis
COMMELINACEAE
COMPOSITAE .
Conferva ambigua
fuscopurpurea
granulosa

178, 179, 180, 227

ae
50

64

59

55

54

58

96
107
100

99
99
100
abd

103
227

227
233-277
181
173-232
180, 183
205

97
e192
179,192
179,194
187
117-118
111

96

2 As
179, 203
100

116

116

103

187

208

199

Conferva—contd.

vadiosa

vupestris

tortuosa
CONNARACEAE
Convallaria fruticosa
CONVOLVULACEAE .
Convolvulus nil
Corallina goughensis

officinalis
CORALLINACEAE
Cordyline fruticosa
Corymborkis corymbosa .

welwitschit .
CRASSULACEAE
CRUCIFERAE .
Cucurbita lagenaria

sicevavia
CUCURBITACEAB
CYANOPHYTA ;
Cyathula prostrata
Cymbopogon citratus
CYPERACEAE . ;
Cyperus pedunculatus

Darea belangeri
Datura candida
DELESSERIACEAE
Dermatolithon nodulosum
Desmodium mauritianum
ramosissimum
Dianthella
compressa .
Dianthera Bey iicillnts
Dianthus sect. Kohlrauschia
sect. Pseudotunica
sect. Tunica ;
subgen. Proliferastrum
diminutus .
filiformis
glumaceus .
nanteutlit
obcordatus .
prolifer
vecticaulis .
saxifvagus .
velutinus
Dichrocephala bicolor
integrifolia
Dicliptera umbellata
verticillata
DILLENIACEAE :
Dimeregramma baldjickii
Dioscorea alata
cayenensis .
dumetorum
DIOSCOREACEAE
Diplocyclos palmatus
Diporochna quintasii

281

192
194
189

98
116
105
106

179, 213

178, 179, 180,

178, 179, 180

129,

211

282 INDEX
Discoclaoxylon occidentale var. pubescens . 112 Gypsophila—contd.
pubescens 112 glomerata .
Dolichos unguiculatus : 99 gvaminea
Durvillaea antarctica 178, 206 haynaldiana
utilis : 206, 207 illyrica
DURVILLAEACEAE . 206 multicaulis
ochroleuca .
EcTOCARPACEAE 197 pachygona
Ectocarpus globifer . 197 vigida
granulosus . 199 saxifraga
mitchelliae . 201 var. glomerata .
ViVESCENS 201 stricta
Eleutheranthera ruderalis 103 thessala
Enteromorpha africana a Od velutina
bulbosa 179, 180, 184
gunniana 183 Hedyotis herbacea
hookeriana . 184 Herposiphonia paniculata
micrococca . 183 Heywood, V. H.
minima 183 Hildenbrandia
nana : 2 183 Hippia integrifolia .
Entophysalis conferta 179, 180, 181 Hymenocallis littoralis
deusta : 179, 181 Hypolytrum grande
Epipogium roseum . 114
Eriosema glomeratum 99 Imperatia
Euphorbia pulcherrima . 111 piliformis
EUPHORBIACEAE LEY lpomoea alba
Exacum quinquenervium 105 digitata
Exell, A.W. ; 93-118 involucrata
Exell, A. W., and C. A. Stace : 1-46 mauritiana
nil 3 :
Feldmannia globifera 178, 197 Ividaea augustinae .
Fiedleria 129 boryana
illyrica 133 ciliata
Fucus antarcticus 206 cordata
pyriferus 205 var. ciliolata
crispata
GELIDIACEAE Fe ZLO heterococca .
Gelidium regulare . 178,210 laminarioides
sp. 178 micans var. ciliolata
GENTIANACEAE 105 undulosa
Geophila neurodictyon 102 Iridophycus boryanum
Gerardia sessiliflora o, A107 ciliatum :
Giffordia granulosa 178, 199 crispatum .
mitchelliae 180, 201 laminarioides
GIGARTINACEAE 218 undulosum
Gossypium barbadense ‘ 97. Txora coccinea
Gough Island, Marine algae of 173-232
GRAMINEAE : : 117 Jatropha multifida
Gymnosporia annobonensis 97 Justicia umbellata .
Gypsophila sect. Petrorhagia 129
sect. Tunicastrum 132 Kalanchoe crenata
subgen. Tunica 129 Kohlrauschia
alpina 145 diminuta
arenicola 151 glumacea
armerioides 139 nanteutlit
compressa . 136 obcordata
cretica 141, 142 prolifera
dianthoides 158 saxifraga
fasciculata . 150 sibthor pit
gasparrinii 154 velutina

171
155
134
133
151
138
142
154
151
154
145
156
166

; 103
179, 180, 228
119-172

179

103

115

117

129
151

105

106

105

106

106

222

219

222

218

222

222

1 2io

179, 180, 218
eee

"78, 179, 180, 222
219

223

223

218

223

102

112
108

99
129, 160
161
169
164
169
161
151
158
166

LABIATAE
Lachnopylis angolensis
annobonensis
mannii
thomensis
Lagenaria breviflora
siceraria
LAURACEAE
LEGUMINOSAE
Leonotis nepetifolia
LESSONIACEAE
Lewis, J.
LILIACEAE ;
Limodorum voseum
Lindernia diffusa
Liparis welwitschii

Lithothamnion muelleri forma neglectum d
ae 179, 180, 217

neglectum .
LOGANIACEAE
Lola tortuosa .
Lophosiphonia scopulorum
Lophurella sp.
LORANTHACEAE
Luffa aegyptiaca ;
Lychaete novae-zelandiae .
Lycopersicon esculentum
Lyngbya confervoides

Macrocystis pyrifera
MALVACEAE .
Mangifera indica
Marine algae of Gough Island .
Maytenus annobonensis
Megaclinium melanorrhachis
Melampodium ruderale
Melasma indicum var. monticola
MELASTOMATACEAE
MELIACEAE
MELOBESIEAE
Melothria capillacea

cordifolia

gilletia

minutiflora :
Microcladia alternata
Microneis hungarica
Mikania cordata

scandens

sp.
MorRACEAE
Mucuna sloanci

urvens

Nasturtium sinapis

Navicula aqueductae
bacilliformis
caucasica
crucicula
jimbriata
fusticulus

INDEX

108
104

205
104-105
116
114
107
114
217

104
oO
179, 231
179, 229
110

101

187

. 106
179, 182

178,205
97

98
173-232
97
115

103

107

100

O77,

178

101

10L

101

Pe ey-
179,227

Navicula—contd.
granum
harstenit
laevissima .
laevissima .
mutata ,
pseudopupula
var. aqueductae
pupula
vaszaryt
wittrockit
Nicotiana tabacum
Nitophyllum sp.
Nuxia angolensis
annobonensis
congesta é
var. thomensis
mannit
NYCTAGINACEAE

Ochna cf. gilgiana
membranacea
quintasit
OCHNACEAE .
Ocimum basilicum .
Odontidium baldjickit
Oldenlandia herbacea
Oplismenus hirtellus
ORCHIDACEAE :
Oscillatoria nigroviridis .
OSCILLATORIACEAE

Palmella conferta

Pancratium littorale

Panicum brevifolium

Paspalum commersonii
conjugatum

Pennisetum polystachion

Peperomia annobonensis .
hygrophila

Peponia vogelit

Peponium bracteatum
vogelii

Persea americana

Petrorhagia, A revision of the genus ‘

Petrorhagia

sect. Dianthella_
sect. Kohlrauschia
sect. Petrorhagia
sect. Pseudogypsophila
sect, Pseudotunica
subsect. Creticae
subsect. Illyricae
subsect. Saxifragae
subsect. Thessalae
alpina

subsp. olympica
arabica
armerioides

var. laxa

179, 182

119-172

284

Petrorhagia—conld.
candica
cretica
cyrenaica
dianthoides
fasciculata
glomerata
glumacea :
var. obcordata
graminea
hispidula .
illyrica :
subsp. angustifolia
subsp. haynaldiana
subsp. taygetea
kennedyae
lycica
macra
multicaulis
nanteuilii
ochroleuca
pamphylica
peroninii
phthiotica
prolifera .
rhiphaea .
rigida
saxifraga
var. gasparrinii
var. Slomerata
thessala
velutina
PHAEOPHYTA
Phaulopsis micrantha

Phormidium submembranaceum

Phyllanthus reticulatus .
Physalis angulata .
micrantha .
minima
PIPERACEAE .
Pladera decussata
Platylepis glandulosa
Plectonema calothrichoides
Pleurostauron sagitta
Plumeria rubra
Polyscias guilfoylei
Polysiphonia boergesenii
howei
scopulorum
Spies
villum
Polystachya ridleyi
Porphyra tristanensis
Portulaca oleracea .
PORTULACACEAE
Principina grandis .
Psychotria neurodictyon
PUNCTARIACEAE
Punica granatum
PUNICACEAE .

179,

179,

179,

179, 180,

178, 179,.

INDEX

141
142
158
158
150
1 By fa
169
169
155
145
133
136
134
137
147
142
158
151
164

Ralfsia .
Rama dobaciaelaniia
Ramatuella
argentea
crispialata .
var. obtusa
latifolia
maguirei
obtusa
virens ,
Remirea maritima .
pedunculata 3
Rhaphoneis baldjickii
fluminensis
lorenziana .
scutellum ;
Rhizoclonium africanum
ambiguum
hookeri
riparium
subsp. ? validum
var. validum
tortuosum
RHIZOPHORACEAE . ;
Rhodoglossum revolutum
RHODOMELACEAE
RHODOPHYTA
Rhynchosia glomerata
Ricinus communis .
RIVULARIACEAE
Rorippa indica
sinapis
Ross, R.
RUBIACEAE
RUTACEAE

Sansevieria metallica
Saponaria cretica
illyvica
Schizachyrium brevifolium
Schizostauron
andicola
brasiliense
crucicula
Simbriatum
karstenit
lindigianum
ovatum
veichardtianum
vrhombicum
tatricum
verbanium
SCROPHULARIACEAE
Scytosiphon lomentaria .
Sellaphora pupula .
Sida rhombifolia
Silene angustifolia .
Sisymbrium indicum
Sledge, W. A
SOLANACEAE .

187

179, 180, 187
a ey)

179, 189

189

189

189

. 100

179, 180, 225
179, 227

208

99

113

182

96

107

: 96
233-277
106

Solanum dasyphyllum
duplosinuatum
nigyrum
nodiflorum

Solenia bulbosa

Sphacelaria furcigera

SPHACELARIACEAE .

Stace, C. A. .

Stanfieldiella imperforata

Stauroneis andicola
cornuta
crucicula
grunowtt
harstenit
lindigiana
mervimacensts
ovata
veichardtiana
rectangularis
sagitta
smithii
stefanssonti
tatrica
verbania
wittrockit

Stearn, W. T.

Summerhayes, V. S.

Synedrella nodiflora

Talinum triangulare
Tarachia falcata
furcata ;
var. platyphylla
nitida
Telanthera mbalind
Terminalia catappa
hilariana
macrophylla
obovata
oxycarpa
punctata
suaveolens .
tomentosa
vasivae
Tetracera ainifolia.,
Thamnopteris nidus
Thecacoris annobonae
Trichomanes adiantoides
aethiopicum

Tristemma oreothamnos

Tunica . ;
sect. Dianthella .
sect. Eutunica
sect. Fiedleria .
sect. Gypsophiloides
sect. Kohlrauschia
sect. Leptopleura
sect. Pachypleura
sect. Pleurotunica
sect. Pseudodianthus

INDEX
106 Tunica—contd.
106 sect. Pseudogypsophila
106 sect. Pseudosaponaria
106 sect. Tunicastrum
184 subgen. Imperatia
178, 202 subgen. Leptopleura
202 subgen. es
I-46 alpina
116 angustifolia
84 arabica
72 avenicola
59 argentit
60 ayrmerioides
55 var. laxa
69 bicolor
60 brachypetala
69 ciliata
70 compressa .
87 cretica
92 cypria
92 davaeana
92 dianthoides
87 erecta
87 fasciculata .
. 89 filiformis
106-1 07, IIO-III gasparrinit
II4-115 glumacea
103 var. obcordata . ;
gracilis
97 graminea
261 haynaldiana
267 hispidula
262 illyrica
265 subsp. angustifolia
109 subsp. haynaldiana .
100 var. haynaldiana
12 var. taygetea
32 kennedyae .
52 lycica
24 macra
nf nanteutlit
25 ochroleuca .
19 var. phthiotica
21 olympica
96 orphanidesiana
242 pachygona ;
111 var. hirtituba .
261 pamphylica
267 peroninit
100 phthiotica
129 prolifera’ .
159 subsp. nanteuilii
148 var. nanteuilit .
a ek29 rhiphaea
132, 148 rhodopea
160 vigida
145 saxifraga
132 var. gasparrinit
132 scoparia
148 sibthorpit

134,

134,
Ve Vines BG.

133,

134,

285

145
132
148
129
145
132
145
136
143
I51
139
139
139
I51
142
151
136
142
147
136
158
I51
150
147
154
169
169
158
155
134
145
134
136
134
134
137
147
142
158
164
138
149
147
156
142
143
159
159
149
161
164
164
156
134
154
151
154
136
139

286

Tunica—contd.
stricta
syrviaca
taygetea
thessala
var. cyrenaica .
velutina
xevophila
Turraea glomeruliflora

Ulva intestinalis var. nana
lactuca
sinuosa

ULVACEAE

137, 141

178, 179,

INDEX

145
170

156
158
166
151

97

183
186
203
183

Uncaria africana
Urera cameroonensis
URTICACEAE

VERBENACEAE

Vigna alba
unguiculata

Viscum engleri

Xanthosoma sagittifolium
Zehneria gilletii

scabra
scrobiculata

do getetss pees
bsp aereonns

etetrtiiin

Feely
sees
We
vere
PT ETE ce revere

} +
ae
erepese

Pr aesiat statebatelebesaae:

pehebebe
preees
+

Teh

ahha atek etal

ees
bo ghee haict a1
idverbeerertrer’

cers

yeas eat
Hy
shoe abee whe

x

tre
rere

weyte
s

seenneee!

eeserestt

tats
stelatest

ae ret re
«

ay
thy.
arte

tf
~ye

os

bets
+5

4
siz3!

>

3et
verere

Eirsts
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