5 01, hi?
ANNALS OF THE
SOUTH AFRICAN MUSEUM
VOLUME XL
ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XL
TRUSTEES OF THE SOUTH AFRICAN MUSEUM
Cape Town
1952 — 1956
& PRINTED IN THE UNION OF SOUTH AFRICA BY
THE RUSTICA PRESS. PTY., LTD., WYNBERG, CAPE
LIST OF CONTRIBUTORS
J. Lewis
A note on the rediscovery of Thunberg's ' 'Gladiolus flexuosus'' and a complete description
of the plant (Part i, September 1952) . . . . . . . . . . . . 1
Plantae Novae Africanae (Part 1, September 1952) . . . . . . . . . . 6
Some aspects of the Morphology, Phylogeny and Taxonomy of the South African
Iridaceae (Part 2, February 1954) . . . . . . . . . . . . . . 15
Iridaceae— New Species and Miscellaneous Notes (Part 3, May 1954) . . . . 115
A Revision of the genus Synnotia (Part 4, April 1956) . . . . . . . . 137
m 11967
INDEX TO NEW SPECIES AND NEW NOMENCLATURAL
COMBINATIONS
PAGE
PAGE
ACIDANTHERA
Gladiolus (cont.)
fiexuosa
4
pubescifolius . .
..132
Aristea
pubigerus
.. 132
latifolia
9
symmetranthus (fig.)
122
Babiana
IXIA
cuneifolia
• • 131
Leipoldtii
. . 121
pubescens
.. 131
stolonifera
14
Exohebea
Lapeyrousia
fiexuosa (plate)
4
effurcata (fig.)
12
fraterna
•• 132
Moraea
unguicularis
• • 132
macronyx (fig.)
.. 115
Ferraria
unguicularis . .
.. 132
brevifolia (fig.)
.. 119
foliosa (fig.) . .
.. 117
Neopatersonia
falcata
8
Gladiolus
namaquensis (fig.) .
6
flexuosus
4
fraternus
•• 132
Synnotia
jonquilodorus
124
parviflora (fig.)
140
meridionalis (fig.) . .
.. 127
Roxburghii . .
146
pubescens
.. 131
Urginea
pubescens
132
multifolia (fig.)
9
INDEX TO PART 2
Acidanthera, taxonomically invalid in S. Africa, 16
Actinomorphic flowers, in most Iridoideae, 90
Anaclanthe, belongs in Babianiineae, 109; closely allied to Antholyza, no; doubtfully distinct,
65-66
Anapalina, bracteoles longer than bracts, 65; contrasted with Chasmanthe, 98; leaves and bracts
of, 26-29; pseudo-lamina of, 30-35; placed in Exohebineae (Ixieae), 108
Anomalesia, distinct from Petamenes, 99; includes Kentrosiphon, no
Atiomatheca, a subgenus of Lapeyronsia, 52; allied to Freesia, 108; corm of, 78-79
Antholyza, belongs in Babianiieae, 109; closely allied to Babiana, no; reclassification by
Brown, 16
Antholyza ringens, terminal spike of, 65-66
Ant holy zeae Hutch, rejected, 107
Aristea, a primitive genus of Sisyrinchieae, 56-58; allied to Bobartia, 60; belongs in Sisyrin-
chinae, 107; most primitive genus, 86; nature of underground stem, 68
Aristea Wredowia is Pillansia Templemanni, 49
Babiana, flowers actinomorphic or zygomorphic, 90; free bracteoles present, 42
Babianiineae, proposed as subtribe of Ixieae, 1 09
Baker's classification of family, 1 5
Bentham and Hooker's classification, 15
Bobartia, belongs in Sisyrinchineae (Sisyrinchieae), 107; inflorescence types, 59-61; nature of
underground stem, 68; second most primitive genus, 86
Bobartia macrospatha has pedicel and perianth-tube, 93, 104
Bobartia tubata is B. macrospatha
Bracteoles of Iridaceae, 40-42
Bracts and inflorescences, 39-66; value of in taxonomy of family, 64-66
Bulbs, absent in South African genera, 68
Chasmanthe, distinctions from Petamenes, 97-98; perianth-tube differentation, 95
Chasmanthe aethiopica, leaf morphology of, 33-35
Chasmanthe caffra and C. intermedia belong in Anapalina, 98
Chasmanthe floribunda, corm of, 71
Chasmanthe furcata belongs in Cur tonus, 98
Cipurinae, a subtribe of Irideae, 107
Classifications, systems of Iridaceae, 15-16, 101-in
Cleanthe, not distinct from Aristea, 1 7
Corm-tunic, morphology of, 80
Corrns, morphology of, 66-89
Crocoideae, abolishment of as a taxon, 105-106; an artificial tribe, 89; connation of stamens
1 01 ; short pedicels and long perianth-tube, 93
Crocus belongs in Ixieae, 104
Curtonas, a valid genus, 98
Diel's classification of Iridaceae, 15
Dierama, corm characters of, 68, 70-71, 72
Dietes, a primitive genus, 86-87; nature of underground stem, 68; rhipidia of, 60, 62
Dietes bicolor, inflorescence of, 62-64
Diplarrhena, flowers zygomorphic, 90
INDEX TO PART 2
Evergreen-leaved genera, 18
Exohebea, bracteoles longer than bracts, 65; distinct from Gladiolus, 16; leaves and bracts of,
26-29; pseudo-lamina of, 30-35; should include Tanaosolen, 108-109; stigmas retuse, 101
Exohebea ramosa, stigmas bifid, 101
Exohebineae, proposed as subtribe of Ixieae, 108
Ferraria, connation of stamens, 101; rhipidia of, 60, 62; tuberous 'corm' of, 72-74
Flower types of Iridaceae, 90-101
Flowers and systems of Iridaceae classification, 16, 89-1 1 1
Galaxia, allied to Homeria, 104; belongs in subtribe Cipurinae of Irideae, 107; connation of
stamens, 101; inflorescence characters, 104
Geissorrhiza, basal disc of corm, 80; nature of style branches, 101; perianth tube, 65
Geissorrhiza Patersoniae belongs in Gladiolus, 92
Gladioleae, taxonomically invalid, 1 09-1 10
Gladiolus, flowers actinomorphic or zygomorphic, 90
Gladiolus edulis allied to Geissorrhiza Patersoniae, 92
Gynandriris distinguished from Moraea, 94; first-leaf, position of, 78
Gynandriris apetala is Moraea Cooperi, 94
Gynandriris setifolia, corm of, 75, 77-78
Gynandriris stenocarpa is Moraea Cooperi, 94
Haeckel's conclusions on inflorescence, 39-40
Hesperantha, allied to Schizostylis, 67-68; basal disc of corm, 80; flat-based corms of, 78-81
Heterostyly in Iridaceae, 100
Hexaglottis, dorsiventral leaves of, 35-38; corm of, 74, 75, 77-78
Hexaglottis virgata, has pedicel and perianth-tube, 93; inflorescence and flowers of, 104
Homeria, allied to Galaxia, 104; connation of stamens, 101; dorsi-ventral leaves of, 35-38;
corms of, 74; rhipidia of, 60-62
Homoglossum, allied to Gladiolus, 90
Hutchinson's classification of Iridaceae, 1 6
Hysteranthous species, 18
Inflorescence, morphology of, 39-66; of Lapeyrousia, 52-64
Internode corms, 74
Iridaceae, classifications of, 15-16; primitive genera, 56-64; systems of classification, ioi-iii
Irideae redefined, 106-107
Iridinae, defined and contrasted, 107
Iridoideae, connation of stamens, 101 ; corm characters of, 68; flowers actinomorphic (except in
Diplarrhena) , 90; flowers always pedicellate in South African species, and nearly always
with no tube, 93; leaf-type of, 18; most primitive tribe of family, 87; rhizomatous genera,
Iris pseudacorus, inflorescence of, 62
Ixia, allied to Dierama, 7 1 ; bracts of, 65 ; confused status of, 1 6
Ixia monadelpha, connation of steamens, 101
Ixieae, defined, 106; divided into six subtribes, 107-108
Ixiineae, subtribe of Ixieae, 1 1 1
Ixioideae, bifid or bicarinate prophylls, 40-42, 65; corm characters of, 68-69, 70-71, 74; most
advanced tribe of family, 88; perianth-tube present, 93
Kentrosiphon belongs in Anomalesia, 99, 1 1 o
Klattia, member of Nivenieae, 87
Lapeyrousia, flat-based corms of, 78-81; flowers actinomorphic or zygomorphic, 90; inflores-
cence of, 52-56; style-branches occasionally simple, 55
Lapeyrousia cruenta, corm of, 78
Lapeyrousia ejfurcata, style-branches simple, not bifid, 101
Lapeyrousia Vaupeliana, style-branches simple, not bifid, 101
Leaf-sheath, due to presence of ligule, 26
Leaf- type of Iridaceae, 18
INDEX TO PART 2
Leaves, morphology of, 18-39; venation, 24; pseudo-laminate, 30-35
Ligular origins in Moraea, 84-86
Melasphaerula, perianth-tube, 65, 93
Micranthus, allied to Watsonia, 47
Moraea, dorsiventral leaves of, 35-38; corms of, 74-77; primitive and allied to Dietes, 86-87;
rhipidia of, 60, 62
Moraea Cooperi, a link with genus Iris, 94; inflorescence and flower characters, 104; redescribed,
94; synonyms of, 94
Moraea plumaria, scale leaf of, 84-86
Moraea ramosissima, spinous roots of, 81-84
Nivenieae, a distinct tribe, 87; defined, 106
Pax's classification, 15
Perianth-tube as a taxonomic character, 93-97
Petamenes, distinct from Chasmanthe, 97-98
Petiole, morphology of, 32
Phylogeny of Iridaceae, 103, 11 1
Pillansia, inflorescence correlated with underground parts, 86; nature of underground stem,
68, 70; phyletic position, 88; should be included in Ixieae, 108; taxonomic position,
49-52
Pillansiineae, proposed as subtribe of Ixieae, 108
Plicate leaves, 109
Prophylls, morphology of, 19, 40-42; of Watsonia, 42-47
Protandry in Iridaceae, 100
Pseudo-laminate leaves, 30-35
Radinosiphon, doubtfully distinct from Gladiolus, 1 1 1
Rhizomes, transition to corms, 67-72
Romulea, affinity with Ixioideae, 81; belongs in Ixieae, 104-105; characters of, 105; flat-based
corms of, 78-81; species with long perianth-tube, 93; Syringodea and Crocus comprise a
separate in Ixieae, 105-106
Romulea bulbocodioides, stem leaves of, 80
Romulea rosea, corm of, 80
Romulea triflora, corm of, 79
Romulineae, a new name proposed for Crocoideae, 107
Scale leaf origin in Moraea plumaria, 84-86
Schizostylis, phyletic position, 88; rhizomatous genus, 67, 68; rhizome of, 79
Shrubby genera, 18; origins of, 87
Sisyrinchineae, a subtribe of Irideae, 107
Spinous roots of Moraea ramosissima, 81-84
Stamens, alternate with style-branches in Romulea and Syringodea, 104; connation of in Iri-
dioideae, 101 ; opposite style-branches in Iris, Galaxia and Homeria, 104; value as taxonomic
characters in Iridaceae, 1 00-101
Staminodes in Watsonia Pillansii and W. marginata, 100
Stipules, 38-39
Streptanthera and Spar axis closely allied, 100
Style and style branches as taxonomic characters, 101
Subterranean stems, 66-89
Symmetry of flowers, 90-93
Syringodea belongs in Ixieae, 104
Tanaosolen, bracteoles longer than bracts, 65; distinct from Ixia, 16; not distinct from
Exohebea, 108-109
Tribal relationship, by underground structures, 86-89
Tricuspidate bracts, 29-30, 46
Tritonia, confused status of, 16
Tritonia subgenus Dichone deserves generic status, 100
INDEX TO PART 2
Tritoniopsis, bracteoles longer than bracts, 65; placed in Exohebineae (Ixieae), 108
Tuberous type of underground stem of Iridaceae, 72-74
Tidbaghia (Amaryllid.j, inflorescence type, 58
Underground parts, eight groups of, 85-86; tribal significance of, 86-89
Viscidity in leaves, explanation of, 38
Watsonia, prophylls in cormlets of, 42-47
Watsonia bulbifera, 42-47
Watsonia marginata, flowers nearly regular, 92
Watsoniineae, proposed as subtribe of Ixieae, 107-108
Weimarck's inflorescence studies, 40; inflorescence theory, 64-66
Witsenia, member of Nivenieae, 87; prophyll and leaf development of, 19-26
Wredowia pulchra is Pillansia, 49
S07.<=3
ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XL
PART I, containing: —
i. A Note on the Rediscovery of Thunberg's c Gladiolus flexuosus* and a
Complete Description of the Plant. — By G. J. Lewis, B.A. (With
Plate I.)
2. Plantae Novae Africanae. — By G. J. Lewis, B.A. (With 3 text-figures.)
ISSUED SEPTEMBER 1952 - PRICE is.
printed for the
TRUSTEES OF THE SOUTH AFRICAN MUSEUM, CAPE TOWN
BY THE RUSTICA PRESS LIMITED, COURT ROAD, WYNBERG, CAPE
ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XL
i. A note on the rediscovery of Thunberg" s 'Gladiolus flexuosus' and a complete
description of the plant. By G. J. Lewis.
(With Plate I.)
During the three years of his stay in South Africa, from 1 772 until the beginning
of March 1775, Thunberg sent or took back with him to Sweden a large collec-
tion of dried plants which he collected on the Cape Peninsula and on the three
extensive journeys he made into various parts of the country. Many of these
plants were named and described by Thunberg himself and quite a number by
the younger Linne, to whom it would appear that Thunberg lent his specimens
for description, according to a note by N. E. Brown on the South African
Iridaceae of Thunberg's Herbarium (Journ. Linn. Soc. of London, XLVIII, p. 16
(1928)).
Among the plants described by Linne f. is the one which he named Gladiolus
flexuosus (Suppl. 96 (1781)). There are two sheets of this species, both collected
by Thunberg. One of them, which must be regarded as the type, is in Thun-
berg's Herbarium in the Botanic Garden at Uppsala. The second sheet, which
Linne f. received from Thunberg in 1 79 1 , ten years after the name was published,
is in London, in the Linnean Society's Smithian Herbarium, with which the
younger Linne's Herbarium is incorporated.
As far as is known exactly 177 years elapsed before the flowers of this plant
were again collected in the field, and during that period there has been some
uncertainty about its identity. In 1863 Klatt transferred it to the genus Sphaero-
spora and thirteen years later Baker placed Gladiolus flexuosus L. f. in the genus
Acidanthera, where it remained until 1941. It was found that it was not possible
to uphold the genus Acidanthera in South Africa as it included a mixture of
species belonging to five different genera. (Gladiolus flexuosus, which must now
be transferred to Exohebea, makes a sixth!)
In a revision of the species placed under Acidanthera, which I published in 1941
(Journ. S.A. Bot. vii, p. 28), I made the following remarks about Acidanthera
flexuosa (L. f.) Baker: —
1
VOL. XL. PART I.
)CT9
2 ANNALS OF THE SOUTH AFRICAN MUSEUM
'There is a drawing at Kew of an imperfect specimen without leaves or corm
and with poor flowers. On the sheet N. E. Brown has made the following notes,
which he has published in his paper on the Iridaceae of Thunberg's Herbarium
in Journ. Linn. Soc. xlviii, p. 22 (1928): "This is identical with the type of
Gladiolus flexuosus Linn. f. and equals Acidantheraflexuosa Baker. As the perianth
is irregular I do not understand why it is not a true Gladiolus. There is no speci-
men like it in the Kew Herbarium." He also states, on the drawing only, that
it is allied to Gladiolus pulchellus Klatt.
'Without seeing the type I cannot be certain about this species but in view
of Dr. Brown's remarks it seems best to leave this as Gladiolus flexuosus L. f.
although examination of the type, when this is possible, may prove it to belong
to the genus Engysiphon.''
Since the publication of that paper I have seen the two sheets of Gladiolus
flexuosus L. f. In 1949, during a visit to England and Europe, I examined these
sheets with particular interest as this plant had not, as far as I was aware, been
collected since Thunberg first discovered it. I found that it did not match any
specimens at Kew, as Dr. Brown had already observed, nor was it represented
in any of the South African herbaria or any of the other herbaria I visited, such
as those of the British Museum, the National Natural History Museum in Paris,
or in Geneva. Unfortunately the three or four specimens on Thunberg's sheets
are incomplete, without corms or basal leaves. I noted, however, that it did
not appear to be a Gladiolus but was probably a species of Exohebea, allied to
E. apiculata (Bolus f.) Foster.
Like so many of Thunberg's specimens, no exact locality was given for this
plant and the only clue which might lead to its rediscovery was a note by Thun-
berg that it flowered in February. According to the records of his travels this
should mean that Thunberg collected Gladiolus flexuosus on the Cape Peninsula,
but there can be no doubt that this species does not occur on the Peninsula.
During the compilation of the Flora of the Cape Peninsula this area was fairly
exhaustively searched over a period of several years and although a number of
new species were found and a few of the apparently 'lost' species rediscovered,
Thunberg's Gladiolus flexuosus was not among them.
The following note on Thunberg's methods, which I received from Captain
T. T. Barnard, is therefore of considerable interest : — 'Literally a record of a
plant flowering in February must mean that he collected it in the Cape Penin-
sula. He spent three Februaries in South Africa and all of them at the Cape.
He certainly collected on the Peninsula in Feb. 1773 and possibly, though less
extensively, in Feb. 1774. I doubt if he did any collecting in Feb. 1775 as he
sailed for Batavia on the 2nd March.
'It is most improbable that G. flexuosus was collected in the Peninsula since it
has never been seen there since. We must therefore fall back on the fact, which
I could parallel from other examples, that the flowering times given in his notes
are estimated and not actual. If for example he collected the last few spikes of a
plant early in March it would be labelled Feb. and if he collected the first spike
NOTE ON REDISCOVERY OF GLADIOLUS FLEXUOSUS 3
at the end of January and saw other undeveloped ones coming on it would be
Feb. also. If Thunberg did not collect G.flexuosus in the Peninsula he must have
collected it coming into flower during his journey from Swellendam to Gape
Town between 1 8th January and 26th January 1774. He spent the night of the
25th January at Badenhorst's farm looking up at Babylons Tower.'
It was on 25 January 1951 that this plant was again collected in flower in the
locality where Thunberg spent the night of 25 January 1774, that is, in the
Caledon Division, on the lower northern slopes of Shaws Pass. It was fairly
abundant locally, growing only in very hard, dry soil, a mixture of clay and
small stones. Many of the spikes were still in bud and in others only the one or
two lower flowers on the spikes were open, so there can be little doubt that
Capt. Barnard's surmise is correct, and that Thunberg gave the flowering
month as February as it was only starting to flower towards the end of January.
At the time of flowering there is no sign of basal leaves on the corms and the
cauline leaves and bracts on the flowering spike are dead and golden-brown in
colour. The basal leaves are hysteranthous and had actually been collected
several years before. It was in fact due to the finding of the unusual basal leaves
that this species was rediscovered and can now be placed in its proper genus and
a complete description given of the whole plant.
In August 1937 Captain T. M. Salter collected one leaf on the lower northern
slopes of Shaws Pass (Salter 6853 in Bolus Herb.). There was no flowering spike
and the leaf, with its slender 'petiole' and short obtuse and comparatively broad
'lamina', was not known to me. I suggested that it might be an undescribed
species of Exokebea, and in May 1938 I visited the same locality with Capt.
Salter and made a collection of the corms and leaves of this plant. There were
no flowering spikes on these corms and some of the leaves were obtuse, others
acute.
Owing to work on the Flora of the Cape Peninsula, and to the outbreak of war
and petrol rationing and other reasons, it was not until early in April 1950
that I revisited the same locality and at that time this species was in fruit, with
the basal leaves just beginning to appear. Having seen the type of Gladiolus
flexuosus the previous year I was able to recognize the spikes immediately as
belonging to that plant. Corms were collected and planted at the South African
Museum Herbarium, where they produced green leaves in June 1950 and
flowered in January 1951, and on 25 January, accompanied by Capt. Salter,
I again visited Shaws Pass and collected a large number of flowering spikes.
The solitary basal leaf, which has two prominent veins and slightly undulate
margins and is sometimes obtuse, is unique in the genus, in fact in the family,
although in many respects it is similar to that of two species of Anapalina, a genus
closely allied to Exohebea. In A. triticea and A. Burchellii the basal leaf is also
solitary and hysteranthous and divided into a slender 'petiole' and a linear-
ensiform 'lamina', with either two or three prominent veins and no visible sign
of any secondary veins. The four or five brown cauline leaves are also anoma-
lous in the genus Exohebea but here again they are similar to those of A. triticea
and A. Burchellii.
4 ANNALS OF THE SOUTH AFRICAN MUSEUM
The long bracts and bracteoles and the long perianth-tube are unusual in the
genus but not unique as they occur also in Exohebea apiculata (Bol. f.) Foster
(Hebea apiculata (Bol. f.) L. Bolus; Gladiolus apiculatus Bol. f.), to which this
species is clearly related. In making the new combination in 1939 (in Contribu-
tions from Gray Herb, of Harvard Univ. cxxvii, p. 36) Foster commented on
E. apiculata: — 'Because of its large size this species is anomalous in the genus.
According to the original description, the perianth-tube varies from 2*4 to
6-3 cm. in length, and the inner spathe (which is longer than the outer) from 3 to
4-3 cm. But in all its essential characters it appears, ex descr., to belong to this
group (i.e. Exohebea). It is possible that it is a tetraploid, or even an amphidi-
ploid hybrid, which would account for its large size.'
Except for the final paragraph, I agree with these views expressed by Foster
and consider that the plant originally described as Gladiolus flexuosus is also an
Exohebea and should be placed in that genus next to E. apiculata. In E. apiculata
the basal leaves were described by Bolus f. as linear, narrowed towards the base
and apex, fully developed after the flowering period, and he added that Dr.
Muir remarked that the flowers appear in April before the leaves, these being
fully developed in June. It is evident from these comments that the leaves are
semi-hysteranthous .
A feature observed in the plants of ' Gladiolus flexuosus' grown at the South
African Museum Herbarium is the long delay between the development of the
spike and the appearance of the flowers. The shoots appeared in June and by
October the spike was fully grown. The bracts and bracteoles remained green
for about six or seven weeks, but there was no sign of the flower buds. These
did not appear until December, by which time the bracts and bracteoles had
started to turn brown, and by the time the flowers opened in January they were
completely dry, brown and rigid. In all the specimens collected at the foot of
Shaws Pass the bracts and bracteoles were also brown and dry. In all species of
Exohebea the bracts are more or less brown and usually rather rigid at the time
of flowering.
The corm is characteristic of the genus Exohebea and so are the protandrous
flowers with their narrow unguiculate perianth-lobes, narrow perianth-tube
(although longer than is usual in the genus) and apiculate anthers. The seeds,
which are light and rather sharply and irregularly angled with narrow wings on
the edges, are also typical of the genus. In all these characters it is quite distinct
from Gladiolus and it is therefore proposed to transfer Gladiolus flexuosus L. f. to
Exohebea, with synonomy as follows : —
Exohebea flexuosa (L. f.) comb. nov. Basinym: Gladiolus flexuosus L. f. in Suppl.
96 ( 1 781); Thunb. Diss. Glad. 9, t. 1 (1784); Prodr. 8 (1784). Syn.: Sphaero-
spora flexuosa (L. f.) Klatt in Linnaea xxxii, 726 (1863) ; Acidanthera flexuosa (L. f.)
Baker in Berl. Monat. xix, 15 (1876); Handbk. Irid. 186 (1892); Fl. Cap. vi,
131 (1896).
Description. — Corm subglobose, about 1-5 cm. diam., enclosed in numerous
brown or reddish-brown fibrous tunics prolonged upwards into a fibrous collar
Ann. S. Afr. Mus. Vol. XL.
Plate
GOX
9 3
Exohebea flexuosa (L.f.) Lewis.
NOTE ON REDISCOVERY OF GLADIOLUS FLEXUOSUS 5
1 1— 15 cm. long and 0-8-2 cm. diam. at the top. One or two cormlets are formed
in the axils of leaves on the underground portion of the flowering stem. Stem
simple, flexuose, 15-30 cm. long (above ground), 1-1-5 mm - diam. Basal leaf
solitary, hysteranthous, with a slender petiole 2*5-5 cm - l° n g (above ground)
and a 'lamina' 3-5-5 cm. long, 1-2 cm. wide, acute or obtuse, with 2 prominent
veins and more or less undulate margins ; cauline leaves 4 or 5, brown at time of
flowering, 4-7 cm. long, half sheathing, the upper half with strongly involute
margins, appearing terete. Spike short, compact, 3-5 - flowered. Bract and
bracteoles similar, brown at time of flowering, 2-5-3-5 cm - l° n gj nearly equal in
length or the bracteoles, which are obscurely bicarinate, up to 5 mm. longer
than the bracts. Flowers pale salmon-pink with crimson median lines on the 3
lower lobes only or on all the perianth-lobes; perianth-tube slender, straight or
slightly curved, 3-4-3-8 cm. long; perianth-lobes narrow, unguiculate, obtuse and
apiculate, the upper half becoming recurved as the flowers mature, the upper-
most 2-5-3 cm - l° n g> 5~8 mm. wide near the apex, the others 2-2-3 cm. long,
2-5-4 mm. wide near the apex; 3 lower lobes sometimes united for half their
length, sometimes free almost to the perianth-tube. Stamens inserted about
2 mm. below the top of the perianth-tube ; filaments about 1-5 cm. long; anthers
6 mm. long with an apiculus 1-2 mm. long. Style, when mature, reaching almost
to the top of the perianth-lobes ; style branches 3 mm. long, with flat emarginate
stigmas. Capsule 1-5-2-2 cm. long, about 8 mm. diam.; seeds numerous, about
5 mm. long and 2 mm. wide, with short wings at the base and apex and usually
on the somewhat irregularly angled sides as well.
Without locality, Thunberg (flowers only, Feb.) Type. (Uppsala). Caledon
Division, lower northern slopes of Shaws Pass, 5 to 5 \ miles south of Caledon. —
Basal leaf only, (Aug.) Salter 6853 (Bolus Herb.) ; corms and basal leaves only,
May 1938, Lewis, S.A.M. 52887; Salter 7222 (Bol. Herb.); fruits and immature
basal leaves, April 1950, Lewis, S.A.M. 60708; flowering spikes Jan. 1951,
Lewis, S.A.M. 60708; Salter 9032 (Bol. Herb.).
Description of Plate I.
1. Corm and young leaf (April). 2. Corm and cormlets, with outer fibrous tunics removed.
3. Basal leaf with acute apex (May). 4. Same, with obtuse apex. 5. Inflorescence (Jan.).
6. Bract. 7. Bracteoles. 8. Flower, side view. 9. Flower, front view. 10. Stamen, front
view — x 2. 11. Stamen, back view — x 2. 12. Style branches — x 2. 13. Fruiting spike
(April). 14. Capsule. 15 and 16. Seeds — x 2.
2. Plantae novae Africanae. By G. J. Lewis.
(With text-figures 1-3.)
Neopatersonia namaquensis Lewis. (Liliaceae-Scilleae.)
Bulbus pyriformis, basin versus 1 -2-1-5 cm. diam., in collum 3-3-5 cm. longum
productus, tunicis exterioribus brunneis subcoriaceis. Folia 2, patentia, lanceo-
lata, acuta, 1-2 cm. longa, 2-4 mm. lata, fere conduplicata. Pendunculus gracil-
limus, 5-7 cm. altus (inflorescentia inclusa) . Inflorescentia dense racemosa 1 0-20
floribus. Bracteae virides, parum carnosae, lanceolatae, o-6— 1 cm. longae.
Pedicelli graciles, suberecti vel patentes, 0-7-1-2 cm. longi. Perianthium gamo-
phyllum tubo campanulato brevissimo; segmenta pallide viridia, parum
carnosa, subaequalia, ovato-oblonga, 3-4 mm. longa, circa 2 mm. lata, exte-
riora semi-reflexa, mucrone minute globosa, interiora patentia, obtusa. Stamina
ad 1*5 mm. longa; filamenta deltoidea, partibus inferioribus connatis et basibus
segmentorum adnatis, apicem versus gradatim subulata cernua; antherae
orbiculares, 0-5 mm. longae. Ovarium ovoideum, circa 1 mm. longum; stylus
circa 1-5 mm. longus, ramis 1 mm. longis ad apicem deflexis, stigmatibus retusis.
Capsula ovoidea submembranacea, 5 mm. longa, loculis 2-3 - sementatis; semina
circa 2 mm. longa, testa atra levi nitida.
Hab. Namaqualand: on a stony slope near the roadside between Springbok
and Garies, twenty-seven miles south of Springbok, Lewis, South African Museum
no. 60893 (T)>P e > m S.A.M. Herbarium), fruit 27 July 1950; flowers May 1951.
Bulb pyriform, 1-2-1-5 cm. diam. near the base, produced upwards in a neck
3-3-5 cm. long; outer tunics brown, subcoriaceous. Leaves 2, spreading, lanceo-
late, acute, 1-2 cm. long, 2-4 mm. wide, usually conduplicate. Peduncle very
slender, 5-7 cm. high (including the inflorescence). Inflorescence a fairly dense
raceme with 10-20 flowers. Bracts green, slightly fleshy, lanceolate, o-6-i cm.
long. Pedicels slender, 0-7-1*2 cm. long, suberect or spreading. Perianth gamo-
phyllous, with a very short campanulate tube; segments pale green, slightly
fleshy, subequal, ovate-oblong, 3-4 mm. long, about 2 mm. wide, the outer
semireflexed with a very small obtuse mucro, the inner spreading, obtuse.
Stamens up to 1-5 mm. long; filaments deltoid, with the bases united and adnate
to the bases of the segments, becoming subulate and cernuous towards the apex ;
anthers dorsifixed, orbicular, 0-5 mm. long. Ovary ovoid, about 1 mm. long;
style about 1-5 mm. long; style branches 1 mm. long, deflexed at the apex, with
retuse stigmas. Capsules ovoid, submembranous, 5 mm. long; seeds 2 or 3 in each
carpel, about 2 mm. long, with a smooth black shining testa.
PLANTAE NOVAE AFRICANAE
8
Figure i. Neopatersonia namaquensis Lewis. I. Plant in flower — natural size. 2. Flower, front
view — x 3. 3. Flower, side view — x 2. 4. Bract — x 2. 5. Inner perianth segment — x 2.
6. Outer perianth segment — x 2. 7. Perianth-tube, with segments removed to show the
stamens — x 3. 8. Gynaeceum — x 5. Del. G. J. Lewis.
In 191 2 Schonland described the genus Neopatersonia {Records of the Albany
Museum II, 252), based on a single species, N. uitenhagensis, from the Uitenhage
Division. Although some of the characters of the plant described and figured
here differ to a greater or lesser degree from those of JV. uitenhagensis these are
probably specific variations comparable with those which occur in some other
8 ANNALS OF THE SOUTH AFRICAN MUSEUM
genera in the Liliaceae such as Urginea, Ornithogalum, etc., and there can be little
doubt that this species should be placed in the genus Neopatersonia. The impor-
tant characters which agree with the generic characters of Neopatersonia are : —
(i) The gamophyllous persistent perianth, (ii) The connate subuniseriate
stamens with the wide bases of the filaments united and adnate to the throat of
the very short perianth-tube. The anthers are introrse. (iii) The style has three
short spreading branches deflexed at the apex.
The following differences between this species and JV. uitenhagensis should be
noted : —
The outer tunics of the bulb are subcoriaceous instead of membranous and
the bulb extends up in a long neck. The two very short leaves are spreading
instead of suberect. The peduncle is more slender and the raceme denser. The
bracts in this species are green and slightly fleshy, although in the closely allied
species described below as N.falcata they are membranous and identical with
those of JV. uitenhagensis. The anthers are orbicular instead of oblong and the
capsule ovoid, not subconic and acute. There appear to be only two or three
seeds in each carpel, with a smooth shining testa, whereas in JV. uitenhagensis
the seeds are numerous and the testa verruculose.
The small and rather insignificant plants were in fruit when collected in
Namaqualand at the end of July. Some bulbs were planted at the South
African Museum Herbarium where they flowered in May the following year.
Neopatersonia falcata Lewis. (Liliaceae-Scilleae.)
Bulbus JV. namaquensi similis sed in collum ad 8 cm. longum productus.
Folium i , falcatum dimidio superiore recurvo, conduplicatum, 5-8 cm. longum,
5-6 mm. latum, acutum, parum carnosum. Pedunculus et inflorescentia JV. nama-
quensi similes. Bracteae membranaceae, basi late ovatae margine laceratae, in
dimidio superiore cuspidatae, circa 6 mm. longae. Pedicelli 0-7-1-2 cm. longi.
Flores JV. namaquensi similes sed segmentis 4 mm. longis, exterioribus interiori-
busque mucronulatis. Capsula ovoidea, membranacea, 6 mm. longa, loculis 2-3 -
sementatis; semina ad 3 mm. longa, testa atra levi nitida.
Hab. South West Africa: Halenberg (thirty-three miles east of Luderitzbucht),
on coastal dunes, Dinter 6665 {Type, in S.A.M. Herb, and Bolus Herb.) 30 Aug.
I929-
Bulb like that of JV. namaquensis but with the neck up to 8 cm. long. Leaf 1 ,
falcate with the upper half recurved, conduplicate, 5-8 cm. long, 5-6 mm. wide,
acute, slightly fleshy. Peduncle and inflorescence as in JV. namaquensis. Bracts mem-
branous, ovate at the base with lacerate margins, the upper half cuspidate,
about 6 mm. long. Pedicels 0-7-1-2 cm. long. Flowers like those of JV. namaquensis
but the outer and inner lobes mucronulate, 4 mm. long. Capsules ovoid, mem-
branous, 6 mm. long; seeds 2 or 3 in each carpel, up to 3 mm. long, with a
smooth shining black testa.
This species is closely allied to JV. namaquensis but the bulb has a longer neck
and produces only one leaf which is falcate with the upper half strongly
PLANTAE NOVAE AFRIGANAE 9
recurved, longer and more fleshy than the leaves of JV. namaquensis. The very
slender peduncle and inflorescence are like those of JV. namaquensis but the mem-
branous bracts are like those of JV. uitenhagensis . The capsules and seeds are like
those of JV. namaquensis but, like the flowers, they are slightly larger. The cap-
sules are also more membranous and there are only 2 or perhaps 3 seeds in each
carpel. This plant was described from dried material and in the open capsule
examined only 2 seeds were present in 2 of the carpels but it is possible that
some might already have escaped.
Urginea multifolia Lewis. (Liliaceae-Scilleae.)
Bulbus globosus, 2-5-3-5 cm - diam., tunicis externis membranaceis rubro-
brunneis. Vagina pallida, 0-5-1-4 cm. longa. Folia multa (circa 30-50) filifor-
mia, plus minusve spiraliter torta, 7-8 cm. longa. Pedunculus 7-12 cm. altus,
floribus 10-12 in racemo densi. Pedicelli 4 mm. longi. Bracteae 3-4 mm. longae,
cordatae, in calcarem acutum 3-4 mm. longum productae. Perianthii segmenta
libera alba, externe pallide lilacina, 1-1-2 cm. longa, 4-5 mm. lata. Stamina
erecta, 6-8 mm. longa, antheris filamentis aequilongis. Stylus declinatus quam
staminibus longior, stigmate paululo trilobato.
Hab. Namaqualand: between Garies and Springbok, on a stony slope near
the roadside twenty-seven miles south of Springbok, Lewis, South African Museum
no. 60870 (Type, in S.A.M. Herb.) Aug. 1950; De Kom (Karas Farm), three
miles from Leliefontein in the Khamiesberg, 4,300 ft., Leipoldt 3378 (in Bolus
Herb.) Oct. 1940.
Bulb globose, 2-5-3-5 cm. diam., with membranous reddish-brown outer
tunics. Sheath colourless, 0-5-1-4 cm. long. Leaves numerous (about 30-50),
filiform, more or less spirally twisted, 7-8 cm. long. Peduncle 7-12 cm. high, with
10-12 flowers in a short dense raceme. Pedicels 4 mm. long. Bracts 3-4 mm.
long, cordate, acute, with an acute spur as long as the bract. Perianth segments
free, white, pale mauve outside, 1-1-2 cm. long, 4-5 mm. wide. Stamens erect,
6-8 mm. long; anthers as long as the filaments. Style declinate, longer than the
stamens ; stigma very slightly 3-lobed.
This species is readily distinguished from all others so far described by the
large number of filiform leaves, resembling those of one or two species of
Gethyllis. The plants grow in groups of two or more among the stones. Most of
them became separated when removed from the soil but in the case of those
figured the two bulbs remained more or less united. The bulbs were collected
at the end of July 1950 and planted at the South African Museum Herbarium
where two of them flowered two weeks later. The flowers, which open early in
the morning and close at about 4 p.m., have a strong sweet lemon-scent, rather
like that of Lippia citriodora.
Aristea latifolia Lewis. (Iridaceae-Sisyrinchieae.)
Planta robusta ad 90 cm. alta ; rhizoma breve, lignosum, 1 -5-2 cm. crassum ;
caulis teres basi circa 1 cm. diam. Folia basalia erecta vel suberecta ad 70 cm.
longa plerumque 40-50 cm., 1-5-2-5 cm. lata vel intima interdum 1 cm., acuta,
IO
ANNALS OF THE SOUTH AFRICAN MUSEUM
Figure 2. Urginea multifolia Lewis. 1. Plants — natural size. 2. Bract — x 3. 3. Flower,
front view — natural size. 4. Gynaeceum — x 4. 5. Stamen, front view — x 4. Del. G. J. Lewis.
PLANTAE NOVAE AFRICANAE I I
nervis densis tenuibus aequalibus induta; folia caulina 5-6, infimum ad 40 cm.,
supremum 7-12 cm. longum. Inflorescentia 25-35 cm - longa paniculata, ramis
sat gracilibus erectis vel erecto-patentibus 6-12 cm. longis et ramulis secundariis
1-2 cm. longis; rhipidia binata, 2-4 - flora. Bracteae ramorum inflorescentiae
inferiores ad 9 cm. longae, 2 cm. latae, superiores angustiores submembrana-
ceae, cuspidatae, suprema 1-1-5 cm - longa. Bracteae et bracteolae rhipidiorum
similes, parvae, lanceolatae, acutae vel acuminatae, 7-9 mm. longae, 2-3 mm.
latae, membranaceae, brunneae vel griseo-brunneae. Pedicelli in statu florente
et fructifero 5-7 mm. vel interdum ad 9 mm. longi. Perianthii segmenta subcaeru-
lea vel pallide lilacino-caerulea, 1-2-1-3 cm. longa, 6-8 mm. lata, exteriora
oblonga, interiora obovata et paululo emarginata. Stamina filamentis 5 mm.
longis et antheris 4 mm. longis. Stylus 8 mm. longus, filiformis, stigmate paululo
dilatato ornatus. Capsula oblongo-ovoidea, 8-9 mm. longa, 7 mm. lata, profunde
trilobata, valvis sat crassis coriaceis praedita, multi-seminatis; semina circa
2 mm. longa, irregulariter angulata vel subcuneata.
Hab. Cape Province. Paarl Division: Wemmers Hoek Peak, gully on west
side, 4,000-5,000 ft., Lewis, South African Museum no. 57345 (Type, in S.A.M.
Herb.) 31 Dec. 1944, and Esterhuysen 11267 (in Bolus Herb.); Winterberg,
cliffs on south-east side, 5,000-5,500 ft., Esterhuysen 9621, Dec. 1943; cliffs on
west side of Upper Wellington Sneeuwkop, 5,500 ft., Esterhuysen 12025, Jan.
1 949 ; Caledon Division : in ravine south-east of Somerset Sneeuwkop, under
trees, Esterhuysen 2935, Dec. 1939; Landdrostkop, Stokoe, South African Museum
no. 55658, Nov.-Dec. 1939; Worcester Division: in shade between rocks on
Du Toits Peak, western ridge, 4,000 ft., Esterhuysen 12382, Dec. 1945; in shade
at foot of cliffs on south-west slopes of Du Toits Peak, 4,500 ft., Esterhuysen 16638,
Dec. 1949; at foot of moist cliffs on south-west side of Slanghoek Needle,
4,500-5,000 ft., Esterhuysen 17798; Louwshoek Mountains, Stokoe, South African
Museum no. 58749, Nov. 1946.
Plants robust, up to 90 cm. high; rhizome short, woody, 1-5-2 cm. thick; stem
rounded, about 1 cm. diam at the base. Basal leaves erect or suberect, up to
70 cm. long, usually 40-50 cm., 1-5-2-5 cm. wide or the innermost sometimes
1 cm., acute, with numerous close fine equal veins, rather soft in texture; cauline
leaves 5-6, the lowest up to 40 cm., the uppermost 7-12 cm. long. Inflorescence
25-35 cm. long, paniculate with fairly slender suberect or somewhat spreading
branches 6-12 cm. long and secondary branchlets about 1-2 cm. long; rhipidia
binate, 2-4 - flowered. Lower bracts subtending the branches of the inflores-
cence up to 9 cm. long and 2 cm. wide, the upper narrower, submembranous,
cuspidate, the uppermost 1-1-5 cm - l° n g- Bracts and bracteoles of the rhipidia
alike, small, lanceolate, acute or acuminate, 7-9 mm. long, 2-3 mm. wide,
membranous, brown or greyish-brown. Pedicels in flowering and fruiting stages
5-7 or sometimes up to 9 mm. long. Perianth segments pale blue or pale mauve-
blue, 1-2-1-3 cm. long, 6-8 mm. wide, the outer oblong, the inner obovate,
slightly emarginate, all with a conspicuous median vein and four very fine veins
on either side, twisting spirally as they fade. Stamens with filaments 5 mm. long
I 2 ANNALS OF THE SOUTH AFRICAN MUSEUM
and anthers 4 mm. long. Style 8 mm. long, filiform, with the stigma only very
slightly dilated. Capsules globose-ovoid, 8-9 mm. long and 7 mm. wide, deeply
3-lobed, with somewhat leathery valves; seeds many in each carpel, about 2 mm.
long, irregularly angled or subcuneate.
Moist shady places in kloofs or gullies or at the foot of cliffs near mountain
tops, from 4,000 to 5,500 ft. Flowering season November-January.
Since the publication of Weimarck's monograph of the genus Aristea in 1 940
several collections have been made, all from high altitudes, of the species which
is now described as A. latifolia. It is quite distinct from any of the previously
described species and cannot properly be placed in any of the eight sections into
which Weimarck divided the genus.
The tall robust plants are similar to some of those in the section Racemosae,
especially A. Bakeri and A. capitata, but the many-seeded small capsules, more
or less ovoid and deeply 3-lobed, are quite unlike those of the section Racemosae
in which they are all more or less 3 -winged and have only 2-3 or sometimes up
to 4 seeds in each carpel. The capsules are in fact very similar to those of some
species in the section Euaristea, the largest section in the genus, recorded from
the eastern Cape Province but with the majority of the species confined to Natal,
the Transvaal and Tropical Africa (also two in Madagascar). In Euaristea the
stem is more or less flattened and winged and the pedicels, at least in the flower-
ing stage, are extremely short (? not more than 1 mm.). A. latifolia, with its
terete stem, and with the pedicels of the flowers not less than 5 mm. long, cannot
therefore be included in that section. It appears to be intermediate between the
sections Euaristea and Racemosae and provides a connecting link between these
two groups.
The leaves of A. latifolia are wider than those of any other known species and
the bracts and bracteoles of the flowers are exceptionally narrow.
Thanks are due to the Curator of the Bolus Herbarium for the loan of the
material collected by Miss Esterhuysen. The large number of well-dried speci-
mens available has been of great assistance in drawing up the description of this
species.
Lapeyrousia effurcata Lewis. (Iridaceae-Ixieae.)
L. heterophyllae proxime accedit, sed ramis styli haud furcatis, distinguitur.
Cormus, caulis, folia et inflorescentia L. heterophyllae similes. Cormus basi circa
1-5 cm. diam. Caulis infra terram 2-4 cm. longus. Folium infimum ad 13 cm.
longum. Flores subcaerulea segmentis basin versus cyaneo-maculatis ; perianthii
tubus 1-2-2 cm. longus; segmenta 5-7 mm. longa, 2-3 mm. lata, obtusa vel
subacuta. Stylus e tubo perianthii 2-3 mm. exsertus, ramis brevissimis haud
furcatis.
Hab. Cape Province. Riversdale Division: Elands Drift, near Riversdale, on
limestone koppies, Ferguson, Bolus Herbarium no. 24923 {Type, in Bolus Herb, and
S.A.M. Herb.) Aug. 1946.
PLANTAE NOVAE APRICANAE
13
Figure 3. Lapeyrousia effurcata Lewis. 1. Plant. 2. Flower, front view. 3. Flower, side view
4. Older flower, side view, with perianth lobes removed. 5. Bract. 6. Bracteoles. 7. Style
branches — x 5. All drawings natural size except where stated. Del. G. J. Lewis.
Very closely allied to L. heterophylla (Willd.) Foster (subgenus Sophronia), but
distinguished by its style branches being simple instead of forked.
Corm, stem, leaves and inflorescence like those of L. heterophylla. Corm about
i*5 cm. diam. at the base. Stem below the ground 2-4 cm. long. Lowest leaf
up to 13 cm. long. Flowers pale blue, with a dark blue mark near the base of
each lobe; perianth-tube 1-2-2 cm. long; lobes 5-7 mm. long, 2-3 mm. wide,
obtuse or subacute. Style exserted 2-3 mm. above the perianth-tube, with very
short, unforked branches.
The simple style branches of L. effurcata, the only character which distinguishes
it from L. heterophylla, are rare in Lapeyrousia in which genus the style branches
are characteristically bifid. Simple branches have been observed, although not
previously recorded, in two other species, both in the subgenus Ovieda. One of
these is L. Vaupeliana Dinter, a densely ramified species from South West Africa,
and the other an undescribed species, also from South West Africa, intermediate
between L. Vaupeliana and L. Bainesii. In Dinter's very brief description of the
former there is no mention of the anomalous simple style branches which are
present in all the specimens in the South African Museum Herbarium collected
by Dinter.
14 ANNALS OF THE SOUTH AFRICAN MUSEUM
Thanks are due to the Curator of the Bolus Herbarium for kindly presenting
some of the living plants collected by Mrs. E. Ferguson to the South African
Museum Herbarium.
Ixia stolonifera Lewis. (Iridaceae-Ixieae.)
Cormus parvus, globosus, 5-8 mm. diam., tunicis e fibris tenuibus submembra-
naceis reticulatis compositis. Caulis 21-50 cm. altus, plerumque circa 40 cm.,
interdum ramo brevissimo infra spicam. Vaginae basales 2, membranaceae,
2 '5~4'5 et 5'5 - 8'5 cm. longae; stolones breves ad apicem bulbillis multis exiguis
ex axillis vaginarum et tunicarum cormi orti. Folia 3, suberecta, lanceolata,
acuminata, mollia, 18-30 cm. longa, 0-3-1 cm. lata, semivaginantia. Spica sat
dense 6-12 - flora. Bractea basi lilacina, 7-8 mm. longa, 3 vel 5-nervata, apice
setaceo-tricuspidata ; bracteolae 6-8 mm. longae, ad parum ultra dimidium
coalitae, setaceo-bicuspidatae. Perianthii tubus cylindricus, 2-4 mm. longus;
segmenta ovata, 0-9-1-2 cm. longa, 6-7 mm. lata, pallide griseo-lilacina, basi
purpureo-ornata. Filamenta purpurea, 2 mm. longa; antherae luteae, 5-6 mm.
longae. Stylus tubo aequans ramis purpureis patentibus 3 mm. longis, apice
stigmatifero haud ampliato; ovarium 3 mm. longum, 2 mm. diam.
Hab. Cape Province. Montagu Division: slopes of Kiesiesberg, on a damp
shaded bank near the road, 3,000 ft., Lewis, South African Museum no. 58727
(Type, in S.A.M. Herb.) Sept. 1946.
Corm small, globose, 5-8 mm. diam., with tunics of very fine, almost membra-
nous, reticulate fibres. Stem 21-50 cm. high, usually about 40 cm., sometimes
with a very short branch just below the spike. Basal sheaths 2, membranous,
2-5-4-5 and 5*5-8-5 cm. long; short stolons with numerous very small bulbils
at the expanded apex arise in the axils of the sheaths and corm tunics. Leaves 3,
suberect, lanceolate, acuminate, soft in texture, 18-30 cm. long, 0-3-1 cm. wide,
half sheathing. Spike rather compact, 6-12 - flowered. Bract mauve at the base,
7-8 mm. long, 3 or 5-veined, with 3 setaceous cusps at the apex; bracteoles
6-8 mm. long, united to just above the middle, with two setaceous cusps.
Perianth-tube cylindrical, 2-4 mm. long; lobes ovate, 0-9-1-2 cm. long,
6-7 mm. wide, pale greyish-mauve marked with purple at the base. Filiments
purple, 2 mm. long; anthers yellow, 5-6 mm. long. Style as long as the perianth-
tube with spreading purple branches 3 mm. long, the stigmas not flattened or
dilated; ovary 3 mm. long, 2 mm. diam.
Two features distinguish this species from all others not only in the section
Euixia, in which it should be placed, but in the genus. One is the long setaceous
points of the bracts and bracteoles and the other the unusual structure of the
short slender bulbiferous stolons. The stolons, which arise in the axils of the
scale leaves of the corm and one or both of the basal sheaths, grow in a vertical
direction and at the apex, just below the surface of the ground, they are more or
less flattened, very irregular in shape, and bear dense clusters of very small
bulbils. The bulbils, none of which exceed 2 mm. in length, have a pale sub-
membranous reticulate outer tunic and when this is removed a very hard black
rugulose inner tunic is disclosed, resembling the testa of a seed and quite unlike
the tunics of the mature corms.
The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular
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1 15
2 9
1 19
1 10
1 10
2 12
2 8
3 13
o 1
o 14
3 3
2 2
8 o
2 11 o
i 6 o
3 15 o
Copies may be obtained from —
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except the Geological and Palaeontological parts, which are obtainable from the
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XL
PART II, containing: —
3. Some aspects of the Morphology, Phytogeny and Taxonomy of the South
African Iridaceae. By G. J. Lewis, B.A., Ph.D. (With Plates
II-XXIV.)
ISSUED FEBRUARY 1954 PRICE ijs. 6d.
PRINTED FOR THE
TRUSTEES OF THE SOUTH AFRICAN MUSEUM
BY THE RUSTICA PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE
3. Some Aspects of the Morphology, Phylogeny and Taxonomy of the South African
Iridaceae* By G. J. Lewis, b.a., ph.d.
(With Plates II-XXIV)
Introduction
For the taxonomist the South African members of the Iridaceae present many
problems and a study of this family has shown that in many cases it is impossible
to describe any large group, such as a tribe or genus, without some proviso to
cover the exceptions or apparent anomalies which occur. Even the description
of the family, according to Baker in the Flora Capensis and other botanists,
requires some slight qualification to include a few of the exceptions which occur
in two or three of the genera.
There has been a considerable difference of opinion not only in the conception
of the genera but also of the main groups into which the family has been divided.
In 1883 Bentham and Hooker, in their 'Genera Plantarum', divided the family
into three tribes, Moraeeae, Sisyrinchieae and Ixieae ; the first two of these tribes
they subdivided into several subtribes. Most of the botanists who have worked
on the classification of the Iridaceae since that date have agreed in dividing the
family into three main groups, but have not agreed about the combination of
the genera in two of the groups.
Pax, in the first edition of Engler and Prantl's Pflanzenfamilien (1889), raised
one of Bentham and Hooker's subtribes of the Sisyrinchieae, Croceae, to subfamily
status, Crocoideae, and combined the remaining subtribes of Sisyrinchieae
with those of Moraeeae, treating this also as a subfamily to which he gave the
name Iridoideae. This subfamily he subdivided into several tribes and subtribes.
The third group, corresponding with Bentham and Hooker's Ixieae, he also
raised to subfamily status and divided into three tribes.
Baker's arrangement in the Flora Capensis is the same as that of Bentham and
Hooker, except that he treated the tribes as suborders and the subtribes as
tribes. Diels, in the second edition of Engler and Prantl's Pflanzenfamilien
(1930), also followed Bentham and Hooker's arrangement, treating the three
main groups as tribes. The only alteration he made was to change the order
in which the tribes were placed by putting Sisyrinchieae first, Ixieae second and
Moraeeae third.
* Thesis approved by the University of Cape Town for the degree of Ph.D., 1952.
VOL. XL. PART 2.
1 6 ANNALS OF THE SOUTH AFRICAN MUSEUM
Rendle, in his Classification of Flowering Plants, and Mario th, in The Flora of
South Africa, both followed Pax's arrangement. A very different arrangement is
that of Hutchinson, who, in his Families of Flowering Plants, published in 1934,
divided the family into eleven tribes of equal status. In 1940 Weimarck estab-
lished the tribe JVivenieae ( Niveniineae, apud Weimarck) , thus bringing the number
of tribes up to twelve.
In view of the various discrepancies that exist with regard to the classification
of this family it is clear that the characters which have been relied on for
differentiation cannot be entirely satisfactory. A distinct tendency in the past
to attach too much importance to the morphology of the flower alone, while
other important characters were overlooked or neglected, has led to considerable
confusion. A few examples which illustrate the fallacy of relying on floral
characters alone for grouping the species together are the following:
Antholyza. In 1932 N. E. Brown pointed out the confusion that had resulted
from a wrong conception of Linne's genus Antholyza, and that the genus,
according to Baker in the Flora Capensis, comprised members of nine different
genera, having in common their conspicuous red or red-and-orange flowers with
the perianth tube somewhat abruptly constricted at or below the middle. In
1 94 1 E. P. Phillips grouped four of Brown's genera together and so reduced the
number to six.
Acidanthera. In 1941, after a critical examination of the South African species
of Acidanthera, I found that it was not possible to uphold this genus in South
Africa as the fourteen species placed under Acidanthera in the Flora Capensis
belonged to five or six different genera, the only feature common to them all
being the comparatively long and slender perianth tube.
Ixia. As arranged in the Flora Capensis this included members of the genera
Gladiolus, Dierama, Tritonia (all these placed in Ixia on account of their actino-
morphic flowers), and the monotypic genus Tanaosolen, the last placed here on
account of a superficial resemblance of the flowers to those of the section Hyalis,
but actually a very distinct plant.
The distinct genus Exohebea was treated as a subgenus of Gladiolus from which
it differs in its corm, leaves, stem, bracts, fruits and seeds, as well as floral
characters, such as the marked protandry of the flowers.
Tritonia. This genus, like many others, is badly in need of revision, and as it
stands at present it includes two distinct genera. Species belonging to five other
genera have already been removed from Tritonia.
It is obvious that the structure of the flower alone does not always provide
sufficient characters for differentiation and that it is necessary to take into
consideration the other morphological members of the whole plant. After
working on the taxonomy of this family over a period of several years it became
increasingly obvious that the only way to arrive at a satisfactory solution to some
of the problems was by means of a morphological study of the entire plant and
by taking into consideration the various correlated morphological characters
of the vegetative and floral organs. This undoubtedly provides more definite
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 1 7
indications of phylogeny than relying merely on floral characters, and it is in
fact the only way to arrive at the true affinities of the genera and tribes and to
place with any degree of certainty the doubtful or borderline species which
appear in some cases to connect the genera.
It also became apparent that although a certain amount of work has been
done on the morphology of this family, a great deal still remained to be done.
Arber has described and discussed many of the different types of leaf which occur
in the Iridaceae. Scott and Brebner, and to a greater extent Adamson, described
the anatomy and secondary thickening of some of the shrubby genera. Ingeborg
Haeckel worked on the inflorescences of some of the genera and Pax mainly on
the morphology of the flowers.
Marloth, who spent so much time studying and making observations in the
field, made many interesting and helpful remarks about the morphology of
some of the genera, many of which are illustrated, and another South African
botanist who has studied this family in the field and by her comments has added
much to our knowledge is L. Bolus. Two botanists outside South Africa who,
during recent years, have done some valuable work on three genera of the
South African Iridaceae, namely Weimarck, in his monograph of Aristea and
revision ofJVivenia, and Foster, in his revision of Geissorrhiza, have both discussed
some of the morphological features of these three genera, especially the former,
who discussed in considerable detail some of the different types of inflorescence
which occur in Aristea, Nivenia and some allied genera. Weimarck included with
Aristea the plant which was formerly treated as a separate monotypic genus,
Cleanthe, which was separated from Aristea only on account of the difference in
the size of the inner and outer perianth segments.
It was found that the works mentioned above did not offer any explanation
for some of the structures observed in certain genera of the South African
Iridaceae, and a few of these morphological features which I have examined
are discussed in the following paper. Various processes of reduction and
adnation suggest an explanation for some of the anomalous features. In making
a morphological study of some members of this family the taxonomist is
confronted by one or two problems concerning which the morphologists,
although they have propounded various theories, have not succeeded as yet in
finding any conclusive evidence. These are mentioned in connection with the
various organs concerned.
The paper is divided into four parts. In the first the structure of some of the
leaves and prophylls is discussed. Part II concerns the bracts and inflorescence.
In Part III the subterranean stems are described and discussed. Part IV is
divided into two sections ; the first deals rather briefly with the flowers and one
or two controversial taxonomic groups, and the second with the phylogeny and
classification. A bibliography is given at the end of Part IV.
For the sake of convenience the names Iridoideae, Crocoideae and Ixioideae, given
by Pax to the three main groups variously treated as subfamilies or tribes by the
botanists mentioned above, are used.
1 8 annals of the south african museum
Acknowledgements
The writer wishes to express her sincere gratitude to Prof. R. S. Adamson,
who was in charge of the Department of Botany at the University of Cape
Town when this research was commenced. Although he retired from the
University a few months later, he kindly continued to assist with advice up to
the final preparation of this paper for publication.
Acknowledgement is made to the Fourcade Fund, University of Cape Town,
for a grant towards the cost of publication of this paper.
Part I. The Leaves
i. Introductory remarks
The most disputed organ in the Monocotyledons is almost certainly the leaf,
and several theories concerning the morphological value of this organ have been
advanced. It is not proposed to discuss these various theories in this paper as
this subject has been dealt with in great detail by Arber, who also devoted much
attention to the several different types of leaf which occur in the Iridaceae. My
remarks will be confined to a few features observed in some of the leaves, bracts
and prophylls examined which have not been discussed by Arber.
As a result of my investigations I am inclined to accept Arber's 'phyllode
theory' as applied to the leaves in this family, that is, that the leaf of the Iridaceae
has no true lamina but represents in every case either a petiole and leaf base
or a leaf base alone.
With the exception of the three shrubby genera, Klattia, JVivenia and Witsenia,
and of the genera Aristea, Bobartia, Dietes, Pillansia and Dierama, all of which are
evergreen, the South African Iridaceae are geophytic and in the majority of the
genera the leaves and inflorescences are produced together and wither and
disappear together at the end of the season. In the western Cape Province
'hysteranthous'* species occur in a few of the geophytic genera, such as Gladiolus,
Anapalina and Exohebea. These so-called hysteranthous species flower during the
dry late summer or early autumn months, and in nearly all of them only one
basal leaf is produced and the cauline leaves present at the time of flowering are
very much reduced. Except for a few of the 'hysteranthous' species the leaves
in all cases have a sheathing base and in most of the genera they are of the
so-called isobilateral equitant type. In some species they are terete and in
several genera in the Iridoideae they are dorsiventral. Exceptions to the typical
sheathing base are found in two species of Anapalina and one Exohebea which
have a very distinct, slender, solid 'petiole' and an expanded pseudo-lamina.
These, as will be shown later, are derived from the typical isobilateral equitant
type of leaf.
* The term 'hysteranthous', often used to describe these leaves, is almost certainly incorrect as
the leaves really precede the flowering shoot. They nearly always start to appear as soon
as the plant has flowered, but they arise from an axillary bud near the top of the old corm
which develops into the new corm and bears a terminal flowering shoot after a dormant
period.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 1 9
As the structure of the mature leaves does not reveal any evidence of their
ontogeny it is necessary to look for the earliest stages which can be found in the
development of these organs. For this purpose various bud cones and shoots
were examined and in one or two of those selected some very early stages were
found which seem to offer some explanation for the development and structure
of the leaves. One of these is Witsenia maura, one of the shrubby genera, in which
some of the prophylls could just be differentiated on the axillary vegetative bud
cones on some of the branches. The other plants to which special attention was
paid were two or three species of Anapalina and Exohebea in which rudimentary
cauline leaves are present on the flowering axis. Some of the smallest of these,
just below the spike, gave some indications of development similar to that of the
Witsenia prophylls.
2. The development of the prophylls and leaves of Witsenia maura and the presence of
a laminar rudiment in the early stages.
In looking for indications of the ontogeny of the leaves the prophylls of
Witsenia maura were examined in various stages, from when they could first be
differentiated, at which stage they were little more than i mm. long, until they
reached their full development, when they varied from 6 to 10 mm. in length.
As these prophylls represent the first leaves borne on an axis in which it was
possible to observe one or two very early stages in development they are of
particular interest. The fusion of the prophylls is discussed later in connection
with the bracts and inflorescence in Part II of this paper. There is no doubt that
the prophylls in Witsenia maura, as in most of the genera of the Iridaceae, are
two more or less united foliar members.
The youngest buds of Witsenia maura to be examined consisted only of a bud
cone and two immature prophylls, the latter about 1-25 mm. long, and there
was some evidence of the prophylls being united as well as what appeared to be
a rudimentary lamina separated from the leaf base in each of the prophylls. A
much enlarged abaxial view of one of these very young buds, removed from the
axis, is shown on pi. II, fig. 1. The bud cone is situated between the two
prophylls which are shown in side view in the figure. On the adaxial side the
overlapping margins of the sheaths are united and under the microscope it could
be seen quite clearly near the top that such a fusion had taken place. The united
overlapping margins near the top are indicated by small dotted lines ; the larger
dotted lines represent the free posterior margins of the sheaths which in some
cases overlap near the base but are not united.
At the base, outside each of the prophyll sheaths, a very distinct protuberance
was present, resembling the rudimentary lamina which, according to Eichler,
is first differentiated from the primordial leaf. In the next stage examined it
was found that this protuberance was carried upwards for a short distance,
adnate to the base of the colourless sheath of the prophyll, but after this no
further development takes place in this region. In mature prophylls there is
sometimes a slight indication of the minute dorsal projection which is evident in
the very early stages, but in mature leaves there is no sign of it.
Ann. S. Afr. Mus. Vol. XL.
Plate II
1-4, Witsenia mama Thunb. 1, young prophylls starting to develop, abaxial view, X40; i, bud
cone; ii, sheath; iii, laminar rudiment. 2 and 2a, immature prophylls, X 2. 3, prophylls, side
view, X2|; i, bud cone; ii, sheath; iii, short monofacial tip; iv, laminar rudiment; v, main
lateral vein; vi, median vein; vii, bud. 3a, plan showing position of prophylls and leaf of shoot.
4, diagram of mature prophylls ; i, sheath; ii, monofacial tip; iii, laminar rudiment; iv, main
lateral vein; v, median vein. 5, Klattia partita Baker, part of old stem and leaves; $a, part of
young stem and leaves, Xi[; i, leaf base; ii, absciss layer; 5^, prophylls, adaxial view, X3I;
5c, prophylls, side view, X 3^. 6-7, JVivenia Stokoei (Guthrie) N. E. Brown. 6, prophylls with
rudimentary lamina, side view, X 2^. 7, part of stem and leaves; i, internode of bud adnate to
axis.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDAGEAE 21
On pi. II, fig. 2, is shown a pair of half-developed prophylls, removed from
the axis and more or less flattened. In these the limbs have started to develop
and in the smaller prophyll on the right there is still a very distinct protuberance,
no longer at the base but extending up nearly to the middle of the sheath, while
in the larger prophyll on the left there is a mere trace of it near the base. The
shaded part is green and the remainder of the two sheaths is colourless. In fig. 2<3
is shown another pair of prophylls at a slightly later stage, with a minute protu-
berance near the base of each. A still later stage in the development of the
prophylls is illustrated in fig. 3. At this stage the prophylls are almost fully
grown and the first leaf of the bud above the prophylls had started to develop ;
its position is shown in a diagram in fig. 3a, where the axis is marked with a
cross and the bud, with its united prophylls and first single leaf, is enclosed
within the sheath of the outer subtending leaf. This leaf above the prophylls
was removed from the bud cone which was cut in half, so that the prophylls
in fig. 3 are shown separated and in side view.
The first vascular strands to enter the prophylls from the axis cylinder are the
main lateral ones and in the prophyll on the right in fig. 3 only these main lateral
vascular strands are present. In the larger prophyll on the left the median
strand is also present and the limb extends above the sheath into a very short
monofacial tip, flattened in the vertical plane, in fact the beginning of the
so-called ensiform isobilateral type of leaf which occurs in Witsenia and so many
other members of this family. It can be seen at this stage that the main lateral
veins enter the apex before the median vein which, at the base, within the bud
cone, is attached to an axillary bud. This is the only sign of a bud in the axil
of these prophylls but it does not develop and the prophylls in Witsenia, as in
many of the other genera, are always sterile. In the smaller prophyll on the
right there is a fairly distinct protuberance near the middle and in the one on
the left a slight trace of it. A diagrammatic representation of one of the largest
prophylls found (1 cm. long) is given in fig. 4.
A few prophylls from some herbarium specimens of Klattia partita and JVivenia
Stokoei, two other members of the small group of shrubby South African Iridaceae,
were examined for comparison. In Klattia partita the two sheaths overlap much
further and are almost, or sometimes completely, united, having a single
common apex, and it is not so evident that the prophylls are two separate foliar
members. At the base of each is a projecting portion corresponding with the
protuberances in the Witsenia prophylls. Two views of these are shown on
pi. II, figs. 56 and 5c. In JVivenia Stokoei the prophylls are also very closely united
and on the poscerior side the overlapping margins at the base are united,
forming a closed sheath round the bud cone. In one of those examined the
structure which appears to be the rudimentary lamina had developed as far as
the top of the prophyll, adnate to it for three-quarters of its length but with the
upper quarter free. A side view of this is shown in fig. 6, on pi. II.
These protuberances may actually be traces of a rudimentary lamina. As far
as can be observed in the prophylls, the petiole starts to develop and the laminar
rudiment is carried up for a very short distance but no further development
22 ANNALS OF THE SOUTH AFRICAN MUSEUM
takes place in the region of the laminar rudiment which gradually disappears.
At about the point where the growth in this region has stopped the median
vascular strand diverges out towards the dorsal margin and the growth
continues in the dorsal area which is now laterally flattened and without a
laminar rudiment. If this is correct then it bears out Arber's phyllode theory,
for the structure which continues to develop and forms the limb is petiolar, with
a flat dorsal wing-like region and no laminar rudiment. As has been shown in
the case of the prophylls, the protuberance begins to disappear at an early stage,
and in mature leaves there is no trace of it.
There are two possible interpretations for the sheath which is so conspicuous
in the early stages in the development of the Witsenia prophylls. One is that it
represents the whole of the remainder of the primordial leaf, that it is in fact
the leaf base, and the other that it is a stipular or ligular 'axillary' outgrowth
from the leaf base, and that the petiole develops outside but completely adnate
to this sheath. The position of the protuberance which possibly represents the
laminar rudiment and the development of the prophylls in Witsenia maura, also
the fact that an absciss layer is formed near or at the base of the limb in Witsenia,
Klattia and JVivenia, seem to suggest that the latter is the correct explanation.
This view perhaps appears similar to that maintained by Gluck, who,
according to Arber, regarded the sheaths of the Monocotyledon leaves as
'stipulae adnatae' which have arisen by the fusion of a pair of free stipules with
the base of the petiole; but my hypothesis is not the same, for judging by the
manner in which the petiole starts to develop outside the sheath in the Witsenia
prophylls, the sheath is definitely not a pair of lateral stipules united to the
petiole but a single stipule or ligule of the axillary type which is not uncommon
in the Monocotyledons and is found in a few Dicotyledons as well. In the fully
developed leaves this ligular sheath is entirely or almost entirely enclosed within
and adnate to the amplexicaul base of the petiole.
There seems to be a possibility that the structure of these leaves, in which the
main lateral veins enter the leaves from the axis cylinder before the median
vascular strand, is associated with the partial adnation of the buds to the axis.
In some of the shrubby species it is apparent that the lowest internode of the bud
is to a greater or lesser extent adnate to the axis. In JVivenia Stokoei the buds are
adnate to the axis for a considerable distance above the base of the subtending
leaf, in some specimens as far as the next leaf above. This was most conspicuous
in a particularly robust specimen of JV. Stokoei with longer internodes than are
usual in this species. A portion of this axis is shown on pi. II, fig. 7, with two
of the leaves removed so that the buds can be seen.
The branching of Witsenia maura was examined and in this genus also it is
evident that the first internode of each branch is adnate to the axis. This is
shown diagrammatically on pi. Ill, fig. 1. Here, as in the rhizomes and corms
of the Iridaceae, the branching is sympodial. After the main axis has flowered
the buds at two consecutive nodes near the base of the stem develop and form
lateral branches. Neither Scott and Brebner nor Adamson, in their descriptions
of the anatomy cf some of the shrubby Iridaceae, mentioned this adnation, and
Ann S. Afr. Mus. Vol. XL.
Plate III
1-5, Witsenia maura Thunb. 1, diagram of stem with two branches partly adnate at the base.
2, apex of stem, with lower leaves removed; 2a, lower part of outer leaf shown in fig. 2, X3;
i, internode; ii, node; hi, leaf. 3, part of old stem with bases of two leaves; i, bud; ii, trans-
lucent margin; iii, leaf base; iv, absciss layer. 4, transverse section of internode at top of stem,
X 10; 4a, t. s. of leaf base, X 10; i, median vein; 46, t. s. of base of leaf, X5^; i, meristematic
tissue; ii, sclerenchyma; iii, median vein; iv, secondary vein; 4c, t. s. about 5 mm. above base
of leaf shown in fig. 2, X 10; i, median vein; ii, main lateral veins ; iii, secondary veins. 5, part
of meristematic tissue, X c. 135; i, sclerenchyma; ii, rudimentary vascular strand. 6, Aristea
oligocephala Baker, bracteole.
24 ANNALS OF THE SOUTH AFRICAN MUSEUM
the whole matter requires investigation. Adamson referred to the fact that the
central parts of the stem are formed before the outer parts, and that the laying
down of the outer parts and the thickening of the outer ground tissue takes place
after the complete differentiation of the central portions. This delay in the
formation of the buds and the subsequent elongation of the internodes with the
still undifferentiated bud tissue adnate to the axis, probably due to intercalary
growth commencing at an early stage, is possibly connected with the delay in
the appearance of the median vascular strand which, instead of passing directly
into the outer region of the leaf base, remains adnate at the base to the axillary
bud and only diverges out towards the dorsal margin after the leaf has started
to develop and the main lateral veins have extended up to the apex. The
median vascular strand in the leaves of the Iridaceae is nearly always very small
in comparison with the lateral veins and in some of the dorsiventral leaves there
is no sign of it.
The internodes just below the apex of the stem in Witsenia maura are extremely
short, but in the older part of the stem they are fairly long, considerably longer
than in Klattia or JVivenia. The apex of a stem of W. maura showing part of the
upper leaves and two internodes and nodes (with the lower leaves removed) is
figured on pi. Ill, fig. 2. Fig. 2a is the upper of the two internodes and base of
the outer leaf, enlarged to show the veins in the base of the leaf, the dotted lines
indicating secondary veins. Two longer internodes near the base of the same
stem are shown in fig. 3. In all these internodes there is a narrow, colourless,
and somewhat translucent margin to the axis just below the dorsal region of
the leaves, which extends up for a short distance into the limb before it gradually
disappears (marked with dotted lines in fig. 3), and on the other side of the
elliptical axis is a similar but shorter and narrower wing-like area on the axis
below the node. In the short internode examined at the top of the stem the
outgrowth of the axis below the dorsal margin of the leaf appears to correspond
with the position occupied by the base of the protuberance in the prophylls.
In the longer internodes it is the same, but owing to intercalary growth the basal
portion is decurrent on the axis throughout the whole internode, while the upper
part can be seen to extend up for a short distance on the dorsal margin of the
limb.
Some transverse sections of the stem and bases of young leaves of Witsenia
maura were examined and, as in the case of the prophylls, it was found that the
first vascular strands to leave the axis cylinder and pass out into the base of the
limb are lateral ones. Some of these sections are shown on pi. III. The first,
fig. 4, is a section of an internode in which two lateral bundles are starting to
leave the central cylinder. In fig. 40 two lateral bundles have entered the arms
of the limb which are already separated from the axis, but the remaining
bundles have not yet left the axis cylinder in the region where the leaf base is
still attached, and the median strand is still some distance inside the cylinder.
In the lower half of this section the vascular strands which will enter into the
next leaf above are oblique as they are just leaving the centre of the cylinder
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDAGEAE 25
and curving out to the margins. (This corresponds with the course of the
bundles in Nivenia corymbosa {Aristea corymbosa) described by Scott and Brebner.)
The third section (fig. qb) is through the base of one of the slightly older leaves,
immediately above the node. The median and twelve lateral vascular bundles
which have entered the leaf from the axis are present, also a few traces of
secondary strands and, near the dorsal margin, a slightly larger vascular bundle.
This dorsal bundle is not the median one and has not come from the central
cylinder but arose in the tissue in the wing-like projection of the axis (in this
section it is the triangular region outside the semicircle of main vascular bundles) .
The tissue in this area was found to be meristematic ; a small portion, very much
magnified, is shown in fig. 5. A few strands of sclerenchyma are formed,
especially towards the margins, and also a few rudimentary vascular strands,
one of which, with a distinct xylem vessel, is present in the portion illustrated.
A few of these bundles were slightly larger than the one shown, the largest being
the one near the dorsal margin.
A few similar rather rudimentary bundles are formed in the external wing-
like portion of the leaf base below the two arms of the limb which, as can be seen
in fig. 4<2, are not supplied with vascular bundles from the axis cylinder. The
small rudimentary bundles in the margins are not connected with the main
vascular strands and seem to disappear towards the top of the sheathing part of
the limb where they are apparently replaced by the primary lateral vascular
strands which diverge slightly outwards towards the margins (see figs. 20 and 3
on pi. III).
Shortly above the base the median vein starts to diverge out towards the
dorsal margin. A fourth section of a young Witsenia leaf is illustrated in fig. ^c
on pi. Ill, which shows the median and two of the main lateral veins starting to
diverge outwards and might be compared with the figure of the prophyll on
pi. II, fig. 3, in which the course of the median vascular strand is shown. In
fig. 4.C (pi.. Ill) the shaded parts represent the assimilatory tissue and the fairly
numerous secondary veins are also shown. The outer dorsal meristematic tissue
is starting to disappear and no trace of it was found in sections cut near the
middle of the leaf.
Near the base of the leaf a fairly distinct absciss layer is formed as the leaves
develop. The position of this layer is marked on the leaf in fig. 3. Scott and
Brebner mentioned that there were distinct indications of such a layer in the
leaves of Nivenia {Aristea) corymbosa. In Klattia partita it is still more evident
(see pi. II, figs. 5 and 5*2), and in this plant the whole amplexicaul base extends
up for a short distance above the node, the lateral arms more than the dorsal
region, and it is persistent on the axis after the leaf has fallen, as can be seen in
the lower part of the stem in fig. 5 on pi. II.
The formation of an absciss layer near the base of the limb seems to suggest
that it is at this point that the leaf base ends and the leaf, or rather the petiole,
begins, and lends some support to my hypothesis that the petiole has developed
outside and completely adnate to an axillary ligule. With the exception of some
26 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the dorsiventral leaves, probably nearly all the leaves in the Iridaceae have
developed in a similar manner. In this respect my views are not entirely in
accordance with those of Arber who regarded the sheathing portion in all cases
as the leaf base, whereas if my interpretation of the sheath is correct, then the
whole limb above the absciss layer, which includes the sheathing portion of the
limb, is petiolar, the sheathing base being due to the presence of the ligule.
In the Witsenia prophylls no development takes place in the marginal regions
of the leaf base and only the dorsal part of the limb develops. The development
of the bracts and bracteoles of some species of Aristea is very similar, as a
comparison of the bracteole of Aristea oligocephala, illustrated on plate III, fig. 6,
with the prophylls of Witsenia mama (fig. 2 on pi. II) will show, allowing for the
fact that the Witsenia prophylls are two partly united members whereas the
Aristea bracteole is a single member. The membranous apex of the sheath in
the Aristea bracts and bracteoles is ruptured as the flower-buds develop and the
single axillary sheath thus appears to be two lateral wing-like stipules.
The leaves of JVivenia Stokoei appear to be exactly the same as those of JVivenia
corymbosa. I have not examined either of these in detail but the latter were
described by Scott and Brebner and according to their description they are
very similar to those of Witsenia maura. Scott and Brebner stated that there were
two median bundles lying one behind the other in the same radial plane, and
also referred to the presence of isolated strands of sclerenchyma in the leaf,
especially toward its edges. There can be little doubt that the outer of the two
bundles which they called median corresponds with the outer dorsal bundle in
Witsenia maura. According to them the inner of the two median bundles (i.e.
the true median bundle), on entering the stem from the leaf, turns sharply
upwards, then sharply down again. This is probably due to its attachment to
the base of the bud which in JVivenia corymbosa, as in JV. Stokoei, is carried up
above the base of the leaf.
In JVivenia Stokoei and other species of JVivenia the dorsal region of the leaf base
is separated from the two lateral arms and the limb develops obliquely with the
base of the dorsal part decurrent on the axis (see pi. II, fig. 7). This may have
something to do with the stage at which intercalary growth commences and is
probably connected with the way in which the bud is carried up on the axis.
The winging of the aerial axis in the plane of flattening of the limb which, as
Arber has remarked, is associated with the equitant leaf in some of the Iridaceae,
is very conspicuous in several species of one or two genera, notably Aristea and
Lapeyrousia.
3. The structure of the rudimentary cauline leaves and the bracts of Exohebea and
Anapalina
In the majority of species in the closely related genera Anapalina and Exohebea
there are rarely any lateral branches and the cauline leaves are very much
reduced, so much so in some species that there is only a trace of them, sometimes
less than 1 mm. long. Several of these rudimentary leaves were examined in
Ann. S. Afr. Mus. Vol. XL.
Plate IV
i , diagram of typical herbaceous bract with reduced tricuspidate bract in dotted lines in lower
half. 2, Tritonia scillaris (L.) Baker, membranous tricuspidate bract, x 2. 3, Ixia micrandra Baker,
membranous bract, X i-|. 4, 4a and 46, Anapalina revoluta (Burm.) N. E. Br., rudimentary cauline
leaves, X2; i, laminar rudiment; ii, prophylls. 5, Exohebea Dodii (Lewis) Foster, rudimentary
cauline leaf, adaxial view, X30; i, bud cone; ii, laminar rudiment; iii, prophylls adnate to
sheath of leaf (v) ; iv, bud in axil of prophyll; 5a, same, abaxial view, X 30; i, laminar x'udiment;
ii, prophyll; iii, sheath of leaf; 56, rudimentary leaf, more advanced stage than preceding, X 1 7 ;
i, short monofacial apex; ii, ? vestigial trace of laminar rudiment ; iii, prophyll; ^c, ? vestigial
trace of same laminar rudiment, X c. 65. 6, Anapalina revoluta, bract and bracteole, x i|-; 6a,
bract, xi|; 6b, bracteole,
prophylls adnate to bract and bracteole.
28 ANNALS OF THE SOUTH AFRICAN MUSEUM
specimens of Anapalina revoluta, Exohebea Dodii and one or two other species and
two or three very early stages in development were found which seem to confirm
my observations regarding the development of the prophylls in Witsenia mama,
that is, that a laminar rudiment is differentiated from the leaf primordium at
the beginning but no further development takes place in this region and it very
soon disappears.
In some of the smallest leaves near the top of the stem in Anapalina revoluta
(just below the spike) a minute cusp-like projection was observed between two
somewhat flattened wing-like structures, the whole less than i mm. long. This
first stage is shown in figs. 4 and A r a on pi. IV. In the second stage (shown in
fig. 46) there is a green and slightly fleshy swelling at the base and the central
cusp is brown. It was thought at first that the wing-like structures on either side
of the cusp were probably free lateral stipules which became adnate to the limb
as it developed but a further examination proved that this was not correct.
A rudimentary leaf from a specimen of Exohebea Dodii was removed from the
stem and when examined under a low-powered microscope it was found that
the lateral wing-like structures are the prophylls of the axillary bud and that
they are almost completely adnate to the sheath of the rudimentary subtending
leaf. Two views of the whole structure are shown on pi. IV, the adaxial side in
fig. 5 and the abaxial in fig. $a. A bud cone is situated between the two prophylls
and in the axil of each prophyll a minute bud is present (see fig. 5). Although
the prophylls are united to the sheath of the leaf they are entirely free from each
other and on the adaxial side they are widely separated. Outside the sheath to
which the prophylls are adnate is the small cusp-like protuberance which is
possibly the laminar rudiment, and which corresponds more or less with the
protuberances found at the base of the prophylls in Witsenia maura. The
protuberance at this stage has already turned brown and begun to shrivel. This
can be seen in fig. $a.
At a slightly more advanced stage, in which the petiole had started to develop
and extended above the sheath in a very short monofacial flattened tip, there
was no visible sign of the cusp-like protuberance found in the very early stages,
but in one of the leaves of Exohebea Dodii examined under a low-powered
microscope a minute hair-like structure was observed a short distance below the
apex. The position of this structure is shown in fig. $b on pi. IV, and the minute
projection, very much enlarged, in fig. 5c. Until there has been an opportunity
of examining a great many more of the leaves in these genera it is not possible
to state with any certainty that this is actually a vestigial trace of a rudimentary
lamina which has been differentiated from the leaf primordium but it seems to
be a possible explanation for this unusual hair-like outgrowth in a plant which
is entirely glabrous. The stage of development reached by this particular leaf
is about the same as that of the left-hand prophyll of Witsenia maura shown in
fig. 3 on pi. II, and the position of the minute hair-like projection more or less
corresponds with the position of the projection on the prophylls, which is most
apparent in the prophyll of JVivenia Stokoei shown in fig. 6 on pi. II, in which
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 2Q,
there is a small brown dorsal outgrowth separated from the remainder of the
prophyll.
The bracts and bracteoles of most species of Exohebea and Anapalina, as well
as those of the closely related monotypic genus Tanaosolen, are cartilaginous and
usually brown at the time of flowering. The bracteoles of these genera differ
from those in all the other genera in the Ixioideae in being longer than the bracts,
sometimes, as in Anapalina triticea and A. Burchellii, very considerably so, but
apart from this the bracts and bracteoles in these genera are almost identical.
A feature which can be observed in most of them, for which at first it was
difficult to find an explanation, is the way in which the fairly distinct outer
margins which appear to be stipular are folded in at the base of the bract and
arise between the axis and the bracteole. The bract and bracteole of Anapalina
revoluta are shown on pi. IV; in fig. 6 they are attached to the axis and in figs.
6a and 6b removed and flattened out. The development of the minute cauline
leaves below the spike provides an explanation for the structure of the bracts
and there can be little doubt that, like the cauline leaves, the bracts have
developed adnate to the prophylls of the axillary bud. This will explain the
thick texture of the bracts, and why the margins arise between the axis and the
bracteole, and also offers an explanation for the anomalous bracteoles in these
genera which, as well as being longer than the bracts, differ from those of the
other genera with a spicate inflorescence in not being bifid or bicarinate. In the
genera Anapalina and Exohebea, as well as in the closely allied genera Tanaosolen
and Tritoniopsis, the prophylls are free from each other but united to the bract,
and the bracteole is therefore almost certainly a third foliar member produced
on the reduced lateral shoot, above the two prophylls and below the flower.
Like the bract, this bracteole has also developed adnate to two prophylls. The
origin of the bifid or bicarinate bracteoles in the other genera with a spicate
inflorescence is discussed later in connection with the inflorescence.
4. The tricuspidate bracts
The tricuspidate bracts which are so conspicuous a feature in Tritonia
subgenus Dichone, as well as many species of Ixia and one or two other genera,
all in the Ixioideae, cannot, in my opinion, be regarded as providing evidence
of lateral stipules. As far as it has been possible to ascertain, the tricuspidation
is due to reduction, the upper half of the bract not being developed. As a result
of this the median and two main lateral veins which are derived from the axis
cylinder remain fairly widely separated instead of converging towards the apex.
This is shown in a diagram in fig. 1 on pi. IV. The large member represents
the type of comparatively large, herbaceous bract found in several genera in
the Ixioideae, such as Gladiolus, Watsonia and Babiana, with only the median and
two main lateral veins marked, the other fairly numerous and uniform veins
present in these bracts being omitted. A line is drawn across the middle and in
the lower half, in dotted lines, is indicated the type of reduced bract found in
Tritonia scillaris, T. trinervata and a few allied species. The actual small reduced
30 ANNALS OF THE SOUTH AFRICAN MUSEUM
bract of Tritonia scillaris, which is about 4 mm. long, is shown in fig. 2 on the same
plate, and a bract of Ixia micrandra (5-6 mm. long) in fig. 3. In the slightly larger
bract of the latter, in addition to the median and two lateral veins there are
two more lateral veins present and traces of six secondary veins starting to
develop at the base.
The bracts of Tritonia scillaris and allied species are membranous and colour-
less, those of Ixia micrandra being herbaceous only at the base where the short
intervening secondary veins are present. From this it might perhaps be deduced
that the bracts in Gladiolus, Watsonia, Babiana, etc., represent the base of a
petiole, while the small reduced bracts of Tritonia scillaris, etc., are only the
ligular outgrowth of the leaf base, with the three (or in Ixia micrandra five)
vascular bundles derived from the axis cylinder, the small intervening veins
present at the base of the bract in Ixia micrandra only appearing as the base of the
petiole begins to develop. This seems to correspond more or less with the early
development of the prophylls in Witsenia maura (except for the presence of the
laminar rudiment in Witsenia), and conforms with the hypothesis that the
petiole in these plants, which all have the isobilateral equitant type of leaf, has
developed outside and adnate to a ligular outgrowth from the leaf base. The
bracts of Sparaxis and Synnotia which are scariose but longer than those of Ixia
and Tritonia subgenus Dichone, are probably derived from the same reduced type
of bract but with the veins more strongly developed and projecting up in three
or five acuminate points.
5. The pseudo-lamina of Exohebea Jlexuosa and Anapalina triticea
In addition to the unusual character of the cauline leaves of most species of
Anapalina and Exohebea, the basal leaves of three species in these genera at first
sight appear to differ so markedly in structure from all the other leaves in the
family that they deserve mention. The 'foliated' leaves of the genus Babiana
have been described by Arber, who pointed out that although they appear to
be divided into a fairly conspicuous 'lamina' and 'petiole', they do not really
differ fundamentally from the ensiform type, and the same is true of the leaves
of Anapalina triticea, A. Burchellii and Exohebea Jlexuosa (L.f.) Lewis {Gladiolus
flexuosus L.f.), all of which have an expanded laterally flattened pseudo-lamina
sharply marked off from a solid, slender petiole-like base. A leaf of A. triticea is
illustrated in fig. 1 on pi. V and two leaves of E. jlexuosa in figs. 3 and 30 on
the same plate, the one with an obtuse and the other an acute apex. The leaves
of the latter, with their slender 'petiole', comparatively broad and short pseudo-
lamina with undulate margins and only two prominent veins, and in many
cases an obtuse apex, are unique in the family. In A. triticea and A. Burchellii
there are either two or three or occasionally four prominent veins; only two
were present in the leaf illustrated.
In most species of Anapalina and Exohebea three or four basal leaves are
produced and are present at the time of flowering, but in mature plants of
A. triticea, A. Burchellii and E. jlexuosa only one basal leaf is produced and this is
Ann. S. Afr. Mus. Vol. XL.
Plate V
i, Anapalina triticea (Burm.) N. E. Br., basal leaf, X \. 2, A. revoluta (Burm.) N. E. Br., basal
leaf, X \. 3 and 3a, Exohebea flexuosa (L. f.) Lewis, basal leaves, x |. 4, E. flexuosa, transverse
section of base of petiole, X 35; i, median vein; ii, main lateral veins; iii, external lateral veins;
iv, fibres; v, phloem; vi, xylem; 4a, t. s. of base of pseudo-lamina, X30: i, median vein;
ii, primary lateral veins; iii, secondary veins; iv, transverse lateral vein; 46, t. s. of middle part
of pseudo-lamina about 2 mm. above previous section, X 30. 5, Exohebea lata (L. Bol.) Foster,
t. s. of sheathing base of leaf, X 30.
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
'hysteranthous'. In all except these three species the linear or linear-ensiform
'blade' tapers to a long, narrow sheathing base. The corm in these genera and
in the closely allied genera Tanaosolen and Tritoniopsis is deep-seated, and the
basal leaves therefore arise a few inches below the surface of the ground, as is
also the case in the genus Babiana.
On comparing the anatomy of the leaves of A. triticea, A. Burchellii and E.
jlexuosa with that of some other members of the same genera it was found that
they are very similar, the only significant difference being the disappearance of
the sheath. As a result of this the basal part is solid and the upper laterally
expanded pseudo-lamina develops equally on either side instead of somewhat
obliquely, as is the case when the sheath is present. (Compare the figures of
the leaves of A. triticea and A. revoluta on pi. V, figs, i and 2.) Some plants of
E. Jlexuosa were grown at the South African Museum Herbarium for observation
so this species has been studied in more detail than any of the others. On some
of the young cormlets which had not yet produced an inflorescence there were
two or three very small leaves and signs of a very much reduced sheath in these.
In mature leaves of E. Jlexuosa the slender basal 'petiole' is solid, with no
opening or sheathing region on the ventral side. The disappearance of the
sheath in this species is almost certainly a recent development as is indicated
by the fact that there are still some signs of a sheath in the immature leaves of
young cormlets, and also by the fact that in the 'petiole' of the mature leaves
the very small external lateral veins are still separate. The median and external
veins are very small in comparison with the two main lateral veins on either side.
The most conspicuous feature in the large lateral veins is the mass of scleren-
chymatous tissue which entirely surrounds and encloses the xylem and phloem.
Between the veins is a small amount of ground tissue consisting of rounded cells ;
towards the top of the 'petiole', in the short portion which is above the ground,
these cells contain chlorophyll. A section near the base of the 'petiole' is shown
in fig. 4 on pi. V and for comparison a section of the slender basal sheathing
part of a leaf of Exohebea lata is given in fig. 5. Apart from the narrow opening
of the sheath the basal part of the leaf of E. lata differs from that of E. Jlexuosa
in having three large lateral veins on either side of the median vein, as well as
five or six minor veins.
At the top of the 'petiole' in E. Jlexuosa the rounded cells of the ground tissue
on either side of the main lateral veins become laterally elongated as the small
median and external lateral veins (the latter now united) curve outwards and
give rise to two or three small secondary veins which are connected with each
other and with the median and external lateral veins by small transverse veins.
The two pairs of large main lateral veins become united to form two pairs of
collateral veins, with the xylem of each separated by a band of sclerenchyma.
At the base of the pseudo-lamina the parenchyma cells between these two united
pairs of main lateral veins are still rounded, as is shown in fig. \a on pi. V, but
about 2 mm. above the base they also become laterally elongated as the main
lateral veins diverge outwards. The small secondary veins which arise in this
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 33
middle portion of the pseudo-lamina are connected with the large primary
veins, as well as with each other, by transverse lateral veins.
After curving outwards near the base of the pseudo-lamina the prominent
primary veins remain widely separated, then in some leaves they converge
gradually towards the apex, in which case the leaf is acute, or somewhat abruptly,
in which case the leaf is obtuse. The course of these prominent veins can be
seen in the figures of the two leaves on pi. V, figs. 3 and 3a. The small median
and secondary veins are not marked in these figures as they are only faintly
visible in dried material and are not perceptible in the living state. Fig. 4^ on
the same plate is a somewhat diagrammatic transverse section about 2 mm.
above the base of the pseudo-lamina, showing the main lateral veins starting
to diverge outwards, with two secondary veins present between them. Chloro-
phyll is present in all the laterally elongated parenchyma cells, though it is
more concentrated in the outer layers, the cells of which are slightly smaller
than those in the middle.
A striking feature in these leaves is the large number of small transverse
lateral veins connecting the xylem of the secondary veins with that of the median
and main lateral veins and with each other. Traces of them are evident near
the base of the 'lamina', as can be seen in figs. \a and aJo on pi. V, but above the
base they are very much more numerous, as can be seen in a section of the leaf
on pi. VI, fig. 1. One of the secondary veins is shown in more detail in fig. la
(pi. VI) with a transverse lateral vein connected with the secondary xylem. In
the linear-ensiform leaves of E. lata no transverse veins were found, but they are
present in Anapalina revoluta, although not as numerous as in E. flexuosa. In A.
revoluta some of the very small secondary veins are united and a few of them
are free, as can be seen in fig. 3 on pi. VI. Fig. 2 on the same plate is a transverse
section of a leaf of Exohebea lata.
The large primary veins of all the species of Exohebea and Anapalina examined
are composed of two united collateral vascular bundles with a small band of
sclerenchyma separating the xylem of the two bundles, and a large mass of
sclerenchymatous tissue between the phloem and epidermis and extending in a
narrow band around the bundles. The median and external veins have no
sclerenchymatous tissue, nor have the small secondary veins which consist of
two united concentric bundles, except some of those in A. revoluta in which
species they are sometimes separate. The median vein of E. lata, which is
identical with that of E. flexuosa, is shown in detail in fig. 2a on pi. VI. In their
small size and particularly in the complete absence of sclerenchymatous tissue
the secondary veins in these genera differ from those in most of the other genera
in the Ixioideae where in many cases they are as iarge as the primary veins and
cannot be distinguished from them. For comparison the upper part of a leaf of
Chasmanthe aethiopica is illustrated in fig. 4 on pi. VI. This type of leaf is very
common in the Ixioideae and has what Arber described as a 'pseudo-mid-rib' in
the region of the main lateral veins. A section of part of this leaf is shown in
fig. 4a, in which it can be seen that there is no difference in size between the
Ann. S. Afr. Mi
Vol. XL.
Plate VI
i, Exohebea flexuosa (L. f.) Lewis, transverse section of part of pseudo-lamina about 2 cm. above
base, X 30; i, median vein; ii, primary lateral vein; iii, secondary veins; iv, transverse lateral
veins; 1 a, secondary vein with transverse lateral vein, x c. 135; i, primary xylem; ii, secondary
xylem; iii, phloem. 2, E. lata (L. Bol.) Foster, t. s. of leaf, X30; i, median vein; ii, primary
lateral veins; iii, secondary lateral veins; iv, fibres; v, phloem; vi, xylem; ia, median vein,
X c. 135. 3, Anapalina revoluta (Burm.) N. E. Br., t. s. of part of leaf, X30; numbers i-vi as in
fig. 2; vii, transverse lateral veins; 3a, part of epidermis and cuticle, X c. 135. 4, Chasmanthe
aethiopica (L.) N. E. Br., upper part of leaf; i, pseudo-mid-rib; 4a, t. s. of part of leaf, X30;
i, fibres; ii, phloem; iii, xylem; iv, assimilatory tissue; v, colourless mesophyll in region of
pseudo-mid-rib. 5, 5a and 56, E. flexuosa, abnormal leaves, x£; 5c, transverse section of the
upper part of the leaf shown in fig. 5a, X 30.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDAGEAE 35
numerous veins, all of which are composed of two united collateral vascular
bundles, except those in the region of the pseudo-mid-rib. All the veins have a
large mass of sclerenchymatous tissue outside the phloem, extending out to the
epidermis.
The leaves of the species of Exohebea and Anapalina examined all have a thick
cuticle and in E. lata and A. revoluta the whole surface is covered with minute
papillae, as shown in a transverse section of a small portion of the epidermis of
A. revoluta in fig. %a (pi. VI). In E.jlexuosa similar protuberances occur only on
the dorsal and ventral margins which are slightly horny.
An interesting irregularity was observed in the leaves of some specimens of
E. flexuosa grown at the South African Museum Herbarium which seems to
suggest that the short and broad pseudo-lamina in this species is of comparatively
recent origin. The corms were collected in the Caledon Division when the
leaves had just appeared but were not fully developed. After being transplanted
the leaves continued to grow and attained the average, or slightly more than
average, length, but the upper half which developed after the corms had been
transplanted was very much narrower than the lower half, as the growth
continued only in the region between the large primary veins. In one of the
leaves there was a very narrow border outside each of the primary veins (see
pi. VI, fig. 5^), and in another none at all (see fig. 5). A third leaf which
developed after the corms were moved was linear-ensiform with almost flat
margins (see fig. 5^), and, except that it was considerably shorter, the 'lamina'
of this leaf resembled that of E. lata. It appears therefore that as a result of
disturbance during the growing-period these leaves reverted to the linear-
ensiform type from which they originated. On comparing a transverse section
of the upper abnormal part of the leaf of E. flexuosa illustrated in fig. $a with a
section of a leaf of E. lata it was found that the former differed from the latter
only in having two instead of three pairs of united primary veins. The transverse
lateral veins characteristic of the broad lower part of the leaf were not present
in the narrow upper part. These two sections are shown on pi. VI, figs. $c and 2.
The structure and development of the leaves in Exohebea flexuosa lend further
support to the theory that the pseudo-lamina of these and many other leaves of
the isobilateral equitant type which occur in many genera of the Iridaceae arises
as a winged outgrowth of the petiole.
6. The dorsiventral leaves of Hexaglottis, Homeria and Moraea
The dorsiventral type of leaf which occurs in most species of Moraea and in
the closely related genera Gynandriris, Hexaglottis and Homeria, has been discussed
by Arber and will only be mentioned briefly here. As Arber remarked, the
leaves of many species of Moraea, etc., are prevailingly dorsiventral, but as a
rule they terminate in a solid monofacial apex, which may often be reduced to
a mere trace. This type of leaf Arber regards as a leaf base crowned by a petiole
and there is little doubt that this is correct. In many of the species the growth
continues for a considerable time in the region of the leaf base which sometimes
Ann S. Afr. Mus. Vol. XL.
Plate VII
i, Hexaglottis flexuosa (L. f.) Sweet, upper half of young leaf, X2; i, leaf base; ii, petiole; iii,
? laminar rudiment; \a, transverse section of leaf base, X 17; ib, t. s. of petiole 1 cm. below
apex, X50; i, fibres; ii, phloem; iii, xylem; iv, assimilatory tissue; v, colourless mesophy 11;
ic, t. s. of swollen tip 5 mm. below apex, X 40; i, assimilatory tissue; ii, group of colourless cells;
iii, colourless mesophyll. 2, Moraea neglecta Lewis, stem with inflorescence and lower half of leaf;
i, leaf base; ii, stipules; iii, petiole; iv, viscid patch on stem below inflorescence; 2a, lower part
of same leaf, X i£. 3, Watsonia meriana Mill., cauline leaf with one stipule; 3a, same, x£; 3b,
bract near top of spike with one stipule, X f . 4, W. humilis Mill., bract near top of spike with
indications of two stipules, X f .
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 37
attains a great length, often up to i metre or more in Hexaglottis longifolia,
H. virgata and H. flexuosa, as well as in a few species of Moraea and Homeria.
Although the growth continues in the region of the leaf base, the growth in the
upper petiolar region often stops at a fairly early stage and as a rule this portion
of the leaf turns brown and disappears when the plants are still quite young.
In the young leaves of Hexaglottis flexuosa and one or two species of Moraea
a feature was observed which was not recorded by Arber, that is, a slightly
bulbous swelling in the petiolar region just below the apex. In young leaves of
H. flexuosa varying from about 6 to 1 2 cm. in length the upper more or less
elliptical petiolar region is only about i -5 cm. long and the swelling about 5 mm.
below the apex (see pi. VII, fig. 1). As the leaf base elongates the whole petiolar
tip dies away and in leaves between 15 and 20 cm. long only a brown, shrivelled
tip remains, but even this dead tip disappears as the leaves become longer.
Some sections of the three regions in the young leaves were examined and some-
what diagrammatic sketches of them are shown on pi. VII. The first, near the
middle of the dorsiventral leaf base (fig. id) is similar in structure to some of the
dorsiventral Moraea leaves described by Arber (e.g. M. bituminosa), with two
very small vascular strands in the mid-rib region. As remarked by Arber, a
striking feature of the dorsiventral Moraea leaves is the unusual character of the
mid-rib region which is thinner than the rest of the leaf and has smaller bundles
which are somewhat irregularly placed, or in some cases they are absent.
The second section (fig. 1 b) is of the petiolar region, about 7 or 8 mm. below
the apex, and the third is of the slightly bulbous tip, about 4 or 5 mm. below
the apex (fig. ic). A few differences are apparent in these last two sections. In
the petiolar region the ribbing, which is fairly prominent in the leaf base, is still
evident, though less pronounced, while in the tip, which is nearly cylindrical
with smooth margins, there is no ribbing and the veins in this swollen region are
arranged slightly towards the centre and are less clearly defined than in the
petiolar region where they are well defined and situated near the periphery. In
the swollen tip six small patches of colourless cells are present between the
epidermis and the green assimilatory tissue which are not present in the petiolar
region, where these patches of colourless cells have evidently broken down and
in the position occupied by four of them there are four grooves. In the region
of the leaf base there are six grooves.
It is possible that the tip, which is differentiated from the petiolar region,
might be a laminar rudiment. The position of this laminar rudiment, if it is
correct to interpret it as such, does not at first appear to be compatible with that
of the protuberances which it has been suggested are probably the laminar
rudiments in the prophylls of Witsenia maura, but actually it lends some support
to the theories already discussed regarding the ontogeny of the leaves. If the
dorsiventral leaf of Hexaglottis, Moraea, etc., is an elongated leaf base, then one
would expect to find a laminar rudiment at the apex. In H. flexuosa a short
petiole is interposed between the leaf base and laminar rudiment. On the other
hand, in the Witsenia prophylls the laminar rudiment differentiated from the
38 ANNALS OF THE SOUTH AFRICAN MUSEUM
leaf base is carried up for a short distance by the petiole as it starts to develop,
though it very soon disappears, but the actual leaf base does not develop any
further, as has already been discussed.
In the dorsiventral leaves the petiolar region is solid and cylindrical (fide
Arber) or somewhat elliptical as in Hexaglottis Jiexuosa, and it is very probable
that in these leaves there is no ligule; it is possible that the disappearance of
the ligule is associated with the development of the dorsiventral type of leaf.
The leaf base entirely encircles the axis and at the base forms a short closed
sheath which extends upwards for a short distance.
In Moraea neglecta Lewis the dorsiventral leaf base is comparatively short
(see pi. VII, fig. 2), and the greater part of the leaf consists of a solid
cylindrical petiole.
A suggestion, which has not so far been investigated, but which might perhaps
account for the viscid patches on the axis below the nodes in Moraea viscaria,
M. bituminosa, Bobartia lilacina, B. paniculata, as well as a species of Homeria and
one or two species of Ferraria from tropical Africa, is that an axillary stipule,
which completely encircles the stem as in Polygonum as well as one or two other
Dicotyledons, is formed at an early stage in these plants, but that it becomes
detached from the leaf base and disintegrates or dissolves into a viscid substance
which is carried up on the axis as it develops. In two of the type specimens of
Bobartia lilacina Lewis in the South African Museum Herbarium (No. 56521),
instead of a viscid patch on the stem below the nodes there are in two cases the
remains of very delicate membranous transparent structures which encircle the
stem immediately below the nodes. One is about 1-5 cm. long and the other
3 cm. long and there are distinct traces of veins on them. These structures are
too delicate to remove from the dried specimens and living material is required
for investigation.
7. Stipules
There are no completely free lateral stipules in the South African Iridaceae
but in a few rare cases the external margins of the sheathing part of the limb,
which is sometimes considerably broader than the upper part, become detached
towards the top of the sheath and are prolonged slightly upwards to form what
might be regarded as short stipular outgrowths. This was found in a specimen
of Watsonia Meriana, but occurred only on one side of one of the lower sheaths
(see pi. VII, fig. 3). At first it was thought that this might simply be due to a
rupture, but in a bract near the top of the spike the same one-sided lobing was
repeated (fig. 36). In a bract near the top of a spike of Watsonia humilis two
slight indications of stipular wings were found (pi. VII, fig. 4) and also in the
leaf of a single specimen of Moraea neglecta (pi. VII, figs. 2 and 2a), but these
represent the only indications of lateral stipules which have been observed. As
has been mentioned in connection with the leaf of Witsenia maura, the extremities
of the wide amplexicaul arms of the limb are not supplied with vascular bundles
from the axis cylinder and the rather rudimentary bundles which develop in
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 39
this region of the leaf base seem to disappear at the top of the sheath, so that the
lateral extremities of the sheathing base of the leaf might perhaps in some cases
be regarded as being stipular.
Part II. The Braots and Inflorescence
1 . Introductory remarks and a brief review of previous works on the inflorescence
In his classification of the flowering plants Hutchinson has attached great
importance to the type of inflorescence in the Monocotyledons, even more in
some cases than to the long-accepted differentiation based on the position of the
ovary. He considered that a classification based on the type of inflorescence
resulted in a nearer approximation of allied genera.
This is undoubtedly true of the Iridaceae, for in this large family the various
types of inflorescences which occur are most important characters for distin-
guishing the tribes, and also, in certain genera, especially in the Iridoideae, for
distinguishing species or groups of species. In some of the larger genera in the
Iridoideae, such as Aristea, Moraea, and one or two others, several different types
of inflorescence occur. It is essential therefore to understand how these various
forms have developed, for they provide most important clues to the phylogeny
of the genera and species. An extensive examination of the inflorescences of
many of the South African genera has shown that the great majority of the
various types have evolved along parallel lines, and that very often links which
are missing in one genus may be found in another.
The interpretation of some of the inflorescences of the Iridaceae by various
botanists in the second half of the nineteenth century has, in some cases, been
proved by more recent workers to be not quite correct from the morphological
viewpoint. This is no doubt due to the fact that their conclusions were derived
mainly from the insufficient material available in the Northern hemisphere,
where a comparatively small number of the genera occur, and these all fairly
highly evolved. Most of the more primitive genera, in which the earlier evolu-
tionary stages can be traced, occur in the Southern hemisphere. Of these a fairly
large number exist in South Africa, particularly in the mountain districts of the
Cape Province. The Cape Province is especially favoured, as in addition to
these primitive forms some of the most highly evolved are found in the coastal
region, e.g. Antholyza, Anaclanthe and Anomalesia.
It is only in the past twenty years that a more thorough examination of the
inflorescences of some of the genera of the Southern hemisphere has been made,
and some of the earlier misconceptions have been corrected and a truer picture
presented of what has taken place. The most valuable work in this connection
is that of Ingeborg Haeckel, published in 1931. The main basis of Haeckel's
work was an anatomical examination of the vegetative cones, following their
development from the earliest stages of differentiation until after the morpho-
logical values of the various parts could be determined. By this method she
arrived at the following conclusions:
40 ANNALS OF THE SOUTH AFRICAN MUSEUM
i . That in many cases the buds of flowers or whole inflorescences which
are apparent in the early evolutionary stages do not develop and reach
maturity.
2. That the spathes or bracts in the groups of the family investigated are
lateral and never terminal, as was previously considered to be the case in
the Iridoideae and Crocoideae in contradistinction to the Ixioideae.
3. The most primitive inflorescence is a much-branched one, the more
composite types being derived from this by various processes of reduction.
In 1939 and 1940 Weimarck published the results of his investigations of some
types of inflorescence in Aristea and some allied genera. On one fundamental
point he does not agree with Haeckel. According to Weimarck the simplest
inflorescences are those in the Ixioideae where they are almost always composed
of a single spike or in some cases lateral branches terminating in few-flowered
spikelets are present as well This he regarded as a primitive arrangement and
he tried to show that the more complicated and composite inflorescences were
derived from this type of spike ; that from the single flower is derived the single
rhipidium in such a way that the prophylls become fertile, and that the binate
two- to several-flowered rhipidia typical of the genus Aristea are similarly derived
from twin or binate flowers.
2. The prophylls and bracteoles
Perhaps the most important point to be considered is the presence of the bifid
or bicarinate prophylls in the spicate inflorescence of the Ixioideae. One of
Weimarck's criticisms of Haeckel's work is that she underestimated or was
unaware of the importance of the bracts and prophylls which remain and, in
the majority of cases, indicate that even the more or less simple inflorescence is
often to be regarded as being derived from a more complicated one. The type
of spike in the Iridaceae differs from a typical or simple spike in having these
bracteoles and their significance was not explained either by Haeckel or
Weimarck.
There has been much discussion about the bicarinate, often bifid, prophylls
and bracteoles which are so common in the Iridaceae and are present in many
other Monocotyledons as well, and two opposing theories have been advanced
in explanation. The first is that the prophylls and bracteoles represent a pair
of foliar members which have become fused together, while the second theory,
favoured by many botanists, among them Arber who made such extensive
studies of the morphology of the Monocotyledons, is that the prophylls and
bracteoles are derived from a single leaf, the two-keeled form being due to a
compression between the main axis and the branch or flower-bud. It is argued
by those supporting this theory that if the prophyll or bracteole were the equi-
valent of two leaves whose mid-ribs formed the keels, it would be expected to
reveal its bifoliar nature by subtending two axillary buds. According to Arber,
however, there is no trace of a second bud.
Ann. S. Afr. Mus. Vol. XL.
Plate VIII
lb
i, Babiana stricta (Ait.) Ker, bract and bracteoles, adaxial view, xi|; \a, bracteoles, Xi|;
lb, part of branch with prophylls at base (adaxial view), xi|; i, part of axis; ii, bract; iii,
bracteoles; iv, top of ovary; v, prophylls. 2, B. plicata (L.) Ker, bract; 2a, partly united
bracteoles; 2b, free bracteoles. 3, B. Sprengellii Baker, bract; 3a, united bracteoles. 4, Babiana
species (aff. B. stricta), transverse section of bract, X 16; 4a, t. s. of one of the bracteoles, X 16;
i, hair; ii, epidermis; iii, assimilatory tissue; iv, vascular bundle; v, xylem; vi, membranous
margin.
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
The genus Babiana is the only one of the Iridaceae known to me in which, in
some of the species, each flower has two distinct and entirely free bracteoles.
An examination of the prophylls and bracteoles of all the species of Babiana
convinced me that these are two distinct foliar members which, in the majority
of species in the genus, have become variously fused. Some of these are illustrated
on pi. VIII. In B. stricta and its varieties and allied species there are two
completely free bracteoles, each strongly carinate, with narrow membranous
margins. At the base, on the side next to the axis, the membranous margins of
the bracteoles overlap slightly, as can be seen in fig. i on pi. VIII, which shows
an adaxial view of the bract and bracteoles of the first flower removed from a
spike of B. stricta. In the case of the prophylls of a lateral branch from the same
plant, shown in fig. ib, the margins do not quite meet. In some other species
two stages have been observed; in some members the bracteoles are separate
while in others the membranous margins are fused at the base, as in B. plicata
(see figs. 2<2 and 2b). Various further stages of fusion are found in other species.
In B. Sprengellii, for instance, the fusion is almost complete, only the two keels
and bifid brown membranous tips remaining to indicate their bifoliar origin
(shown in fig. 30 on pi. VIII).
These various stages seem to indicate clearly that this was a case of fusion
having taken place due to juxtaposition and pressure, and not a case of a single
member having become divided into two free members by pressure. It remained
to find some proof in support of this conviction, but none could be found in the
genus Babiana. A great number of plants from several species were examined
but there was no visible sign of any buds in the axils of the prophylls or bracteoles.
Many sections of the single and fused bracteoles were examined but these
produced no evidence of any actual fusion. The free bracteoles are smaller in
size, more strongly keeled and have conspicuous membranous margins, but
otherwise do not differ in any respect from the bract. Transverse sections of a
bract and bracteole of an undescribed species of Babiana, allied to B. stricta, are
shown on pi. VIII, figs. 4 and \a.
It was necessary therefore to turn to some other genus in the hope of finding
some evidence of significance in connection with the bracteoles. With the idea
in mind that the spike was derived by reduction or suppression from a more
densely branched type of inflorescence, the first plant examined was Watsonia
bulbifera, which provided some definite proof of a process of fusion taking place.
3. Fusion of the prophylls in the cormlets of Watsonia bulbifera
Watsonia bulbifera Mathews and L. Bolus is one of the tallest and most robust
members of the genus and a characteristic feature is the large number of aerial
cormlets or bulbils produced in the axils of the upper sheathing leaves and bracts.
As the authors remarked, the habit of producing cormlets is rare in the genus,
occurring only in two or three species, and in none are the cormlets so numerous
as in this. The few capsules that form (usually with one or more cormlets in
the same axil) contain only one or two seeds which are perfect.
Ann. S. Afr. Mus. Vol XL.
Plate IX
Watsonia bulbifera Mathews & L. Bol. i, node near base of inflorescence ; la, same node with
bract removed showing immature 'bulbils'; i, prophylls. 2, pair of partly united bracts sub-
tending two buds, X 6; 2a, same bracts, back view. 3, partly united bracts with two buds, X 6;
3a, same bracts with buds removed. 4, pair of partly united bracts with one bud, X 6; 4a, same
bracts with bud removed; i, free margin of bract on left. 5, pair of united bracts with common
apex, front view, x6; 5a, same bracts, back view; i, free margin. 6, immature 'bulbil', X7;
6a, bract, x6; 6b, united prophylls, x6; 6c, first scale leaf, X 16. 7, prophylls shown at i in
fig. 1 a, X 2 ; i, free margin of prophyll on left.
44 ANNALS OF THE SOUTH AFRICAN MUSEUM
The number of cormlets in the axils of the bracts at the base of the inflorescence,
where more or less erect lateral branches are present, is greatest, up to about
thirty to a bract, decreasing in number towards the top of the spike. The final
three or four bracts of the spike usually subtend flower-buds only, those just
below having one or two cormlets as well as the flower. One of the nodes near
the base of an inflorescence was examined at the beginning of the flowering
season, at which stage the cormlets are present as small immature buds. An
erect lateral branch was developed, next to the axis, and between this and the
subtending bract were about thirty immature cormlets which even at this early
stage had exerted sufficient pressure on the subtending bract to rupture it.
This is illustrated on pi. IX, fig. i. The same node is shown in fig. \a but with
the bract removed. Immature cormlets were also found in the axils of the lower
bracts of the branch in some specimens.
Many of the cormlet buds were found to be subtended by more or less
enveloping pairs of bracts or prophylls in various stages of fusion. Some of the
bracts were only partially united and some of these subtended two separate buds,
while in others, where the fusion was more complete, only one bud had
developed. Some of these stages are shown on pi. IX. In fig. 2 two bracts can
be seen subtending two buds; in this case the bracts are almost free, with only
the margins united. The outer side of the same pair of bracts is shown in fig. 2a.
In fig. 3 two buds are present but the bracts are more closely united, as can be
seen in fig. 30, which represents the same pair of bracts with the buds removed.
Fig. 4 shows a pair of fused bracts with only one bud developed. This was
removed and the two partially united bracts are shown in fig. 40. Here the
inner margin of the smaller bract on the left is free and can be seen as a small
overlapping flap in the centre. Figs. 5 and 50 are the outer and inner sides of
two bracts which are still more closely united, though the overlapping edges of
both are still free and can be seen as flaps on the outer and inner sides. In this
case the two bracts are more or less united in a common apex and only one bud
was present.
Fig. 6 represents a single young cormlet enveloped in its subtending bract
(fig. 6a). Within this but on the opposite side of the bud are two completely
united sterile prophylls (fig. 6b), which look like a single slightly bicarinate
bract. The next foliar member, which encloses the bud and is united around it
at the base, is the first scale leaf which remains attached to the young cormlet.
There is an opening at the side through which the next scale leaf emerges as it
develops (see fig. 6c). When mature cormlets were examined about two months
later (shown on pi. X, fig. 1) the outer bracts and prophylls were membranous,
very thin and brittle, and tended to fall apart and break up when touched, so
that they evidently do not take any further part in the protection of the young
cormlets after they have developed. This function is performed by the first and
four or five succeeding scale leaves, which at this time were brown and more or
less fibrous. These are shown in a longitudinal section of a bulbil in fig. \a on
pi. X.
Ann. S. Afr. Mus. Vol. XL.
Plate X
Watsonia bulb if era Mathews & L. Bol. i, mature cormlets, x § ; i, remains of bract; ii, branch;
\a, longitudinal section of cormlet, x \\. 2, node of first flower of spike with bract removed, X f ;
i, immature cormlets; ii, bracteoles of flower; iii, perianth tube; 2a, bract from same node,
xf; 2b, bracteoles of flower, X ij. 3, node of first flower (two months later) with mature
cormlets, xf; i, capsule. 4, node of second flower of spike, xf; i, internode on lateral bud;
\a, bud with internode, removed from axis, x \\. 5, sixth bud of spike (adaxial view), xf;
50, bracteoles of same bud, with cormlet buds, X f . 6 and 6a, front and back views of bracteoles
of seventh flower, xf. 7, terminal bud, xf; ja, bract, xf; yb, bracteoles, back view, xf.
8 and 8a, top flower-buds of spikes with vestigial tips, X f .
46 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 7 on pi. IX shows the two prophylls of the shoot. They are situated
between the branch and the axis, as can be seen in fig. 10 on the same plate.
The smaller one on the left subtends a bud and is partly united with the larger
one, with the free margin present as a small flap. The larger prophyll is actually
the one which subtends the branch. (The more or less tricuspidate form of this
prophyll is very unusual in the genus Watsonia and is probably due to the
crowded conditions at the node as a result of which only the median and two
main lateral veins are developed and project up in three short cusps. The
tricuspidate bracts of some other genera are discussed in Part I of this paper.)
Evidently some sort of proliferation has occurred in Watsonia bulbifera which
has resulted in the development of those buds which, in the Ixioideae, are almost
invariably suppressed. Thus buds have developed in the axils of many of the
prophylls which are normally sterile. The prophylls shown in fig. 7 are those
of the branch which is subtended by the large outer bract at the node. All the
internodes of this branch have been suppressed and the lateral branch which is
present is not a primary branch of the main axis but a secondary branchlet, the
first branchlet of the primary branch. The second is present as a bud which
develops into a cormlet through the accumulation of food reserves in its abbre-
viated stem.
The next node examined on the same inflorescence was that of the first flower.
When the outer bract was removed two buds were disclosed, one on either side
of the flower. These are shown in fig. 2 on pi. X. The bracteoles of the flower,
which envelop the ovary and base of the perianth tube, are united for about
three-quarters of their length, with free acute tips. When removed from the
flower and opened up, a bud was found between them and the ovary (see pi. X,
fig. 2b). In fig. 3 on the same plate a later stage in development is shown. At
this stage, about two months after flowering, there are three mature cormlets.
There were no fertile seeds in the capsule.
The second flower in the same spike was the same as the first except that in
the bud on the left an internode had developed, producing a rudimentary
branchlet (see pi. X, 4 and 40). The next four bracts subtended flower-buds,
but here no lateral buds were present next to the flower-buds, though in each
case a bud was found between the ovary and the bracteoles. In the sixth a
second very small bud was present (fig. 5a) . The last four bracts of the spike
subtended flower-buds only, with no visible sign of any other buds.
The fusion of the bracteoles of the spike becomes progressively more complete
towards the top of the spike. At the base they have very distinct free tips but
compare figs. 2b and 5a on pi. X, the latter the bracteoles of the sixth flower in
which the apex is very shortly bifid. The inner and outer sides of the bracteoles
of the seventh flower (figs. 6 and 6a) show the minutely bifid apex and towards
the base two shallow pouches. The bracteoles of the terminal flower are com-
pletely united and have only two rather obscure pouches at the base (shown in
fig. yb). In some of the spikes a vestigial tip was present above the terminal
flower. Two of these tips are illustrated in figs. 8 and 8a.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDAGEAE 47
No living material of Watsonia vivipara Mathews and L. Bolus was available
for examination but in this species a similar proliferation has resulted in the
production of numerous cormlets. The development of the cormlets is not quite
the same as in W. bulbifera, and the authors commented on this as follows:
'The name alludes to the "bulbils" which while still on the parent plant put
forth a green leaf projecting for about i cm. beyond the basal sheath of the
"bulbil", suggesting the idea that this leaf may be functioning in order to permit
the "bulbils" to start growth without a preliminary resting-stage. In the large
number of bulbils produced this species resembles W. bulbifera, but this differ-
ence must be noted : the development of the bulbils is concurrent with the
development of the flowers in W. vivipara, whereas in the former their develop-
ment starts when the flowering is nearly, if not quite, over.'
In the allied genus Micranthus a similar proliferation occurs in one of the
species, M. junceus N. E. Br. In this very much smaller plant with its much
denser spike the number of cormlets produced is not as great as in Watsonia
bulbifera, not more than seven to nine being found sometimes in the axils of the
lowest bracts; more often there are solitary cormlets, or from two to five to a
bract near the base of the spike, or, in some specimens, no cormlets are produced.
The prophylls of Watsonia bulbifera provide fairly conclusive evidence of a
fusion taking place between two closely adjacent overlapping members, the
closer union of the two members resulting in the suppression of the lower of the
two axillary buds.
4. Vestigial traces of bracts in Watsonia pyramidata and a hybrid
Some vestigial traces of bracts were found in one or two species of Watsonia,
as well as in Pillansia Templemanni, which furnish further evidence of the spike
having been evolved from a more complex inflorescence through various stages
of reduction.
When collecting fresh flowers of Watsonia pyramidata (Andr.) Stapf (W. rosea
Ker) in the field it was frequently noticed that within the bracts of some of the
specimens was a colourless mucilaginous substance. What appeared to be a
natural hybrid between W. pyramidata and W. Schlechteri, from the top of the
French Hoek Pass, was sent to me and here again the viscid substance was
observed inside some of the bracts subtending the flowers. One inflorescence
was therefore examined in detail.
It was found that the one or two short, erect branches at the base have their
lowest internode partially adnate to the axis (see pi. XI, fig. la). Between these
branches and the axis were the short, united but distinctly bifid prophylls
which are shown in fig. 2 on pi. XI. In the flowering parts of the inflorescence
small and very delicate imperfect structures were found in many cases between
the closely adpressed bracteoles and the axis. Some of these are shown in figs.
3 and 3a on pi. XI. These structures were translucent, pale pink in colour, and
when moistened they immediately dissolved and became colourless and muci-
laginous. Under the microscope they were seen to be only one or two cells in
Ann. S. Afr. Mus. Vol. XL.
Plate XI
1-4, Watsonia hybrid. 1, first node of inflorescence, bract removed, xf; la, second node;
i, internode of branch partly adnate to axis. 2, prophylls of branch in fig. la, X f. 3, 3a and 36,
adaxial views of bract and bracteoles of flower-buds showing vestigial remains of prophylls, x f;
i, bract; ii, bracteoles; iii, remains of prophylls. 4, part of vestigial remains of prophylls,
x c - x ?)^- 5? Watsonia Middlemostii L. Bol., part of spike, xf; i, outer bract; ii, prophylls; iii,
inner bract; iv, bracteoles. 6-1 1, Pillansia Templetnanni (Bak.) L. Bol. 6, branch of inflorescence,
xf; 6a, bract subtending branch, X i£; 6b, prophylls of branch, xij; 6c, prophylls of secon-
dary branchlet, X r£. 7, apex of primary branch with tip developed, x %,. 8, bud with vestigial
traces of inner bracts, x il; 8a, bract of bud, Xi|; 8b, bracteole, Xi|; 8c, first vestigial bract,
Xi|; 8d, second ditto, X 2; 8?, ovary and second vestigial bract, X 2. 9, diagram of bracts and
flower shown in fig. 8. 10 and 1 1, diagrams of two theoretical earlier stages before reduced stage
shown in fig. 9. 12, Nivenia corymbosa (Ker) Bak., diagram of bracts and branches at first node
of inflorescence; X, main axis; Bi-Bv, bracts.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 49
thickness, consisting only of one or two epidermal layers, with elongated cells
and distinct traces of stomata (see pi. XI, fig. 4), resembling the epidermis of
the bracteoles which was examined for comparison. In a bud near the top of
the spike these translucent structures were present between the bracteoles and
the flower as well as between the bracteoles and the axis (see fig. 3^).
These extremely delicate structures are almost certainly the vestigial traces of
prophylls of which only the epidermal layers are partially developed. That this
is possible seems to be borne out by the arrangement of the bracts in a few
specimens of Watsonia Middlemostii L. Bol. This species is closely allied to
W . pyramidata from which it differs in being smaller in all its parts, with narrower
leaves and nearly always an unbranched axis. In specimens collected by
Schlechter at Onrust River, Klein River Mountains (No. 9492), it was observed
that the flowers in the spike were subtended by two bracts and had two pairs
of united bracteoles between the flowers and the axis, the lower pair of the latter
very short and not very noticeable (see pi. XI, fig. 5), whereas the upper ones
are as long as the inner of the two bracts. One of the outer bracts was removed
and an extremely short internode could just be discerned between the outer and
inner bracts. The reduction from a branched inflorescence to a simple spike
has for some reason been arrested in these plants, and the lower of the two pairs
of united members between the flowers and the axis are therefore the prophylls
of the reduced lateral branch, the upper two, which are almost completely
united, being the bracteoles of the flowers. This is comparable with the arrange-
ment in W. bulbifera where the lower prophylls are present on either side of the
lower flowers in the spike, but instead of being sterile and united they are free
and each subtends a bud.
5. Vestigial traces of bracts in the inflorescence of Pillansia Templemanni
From the phylogenetic point of view Pillansia Templemanni (Bak.) L. Bolus
(Tritonia Templemanni Bak.) is of particular interest. This monotypic genus,
which is confined to a small area near the coast in the Galedon Division, is
indeed not easy to place as it is intermediate between the Iridoideae and Ixioideae.
In many respects it is most closely allied to Watsonia and may be regarded as
the ancestral type of that genus, but, as noted by L. Bolus, it differs among
other things in having flowers which are regular and arranged in a 'cymose
panicle'. Baker and L. Bolus evidently were not aware of the name Wredowia
pulchra given to this plant in 1827 by Ecklon, as neither of them cited this name,
which must, however, be regarded as a nomen nudum. (Apart from stating that it
was intermediate between Sisyrinchium and Aristea Ecklon gave no description
of the genus or species.) In 1840 Steudel, in Nomencl. ed. II, 1, 1300, changed
the name to Aristea Wredowia, but in 1 940 Weimarck quite rightly excluded this
pJant from the genus Aristea. Hutchinson placed Pillansia in the tribe Ixieae.
The systematic position of this plant is referred to again in Part IV.
Some vestigial traces of bracts were found in the inflorescence which provide
some evidence of a link between the spicate inflorescence and a more richly
50 ANNALS OF THE SOUTH AFRICAN MUSEUM
ramified one such as occurs in some species of Nivenia and Aristea. L. Bolus
described the inflorescence as follows : 'Flowers solitary, terminal on the primary,
secondary, tertiary and quaternary branches, forming a somewhat lax panicle
25-40 cm. in length, with ascending or suberect branches 4-9 cm. long.'
In living material from the Palmiet River Valley which was examined there
were fewer branchlets than in the inflorescence described above. A primary
branch is shown in fig. 6 on pi. XI. This is subtended by a short bract (fig. 6a)
and between the branch and axis are a pair of united prophylls, one slightly
longer than the other (fig. 6b). Similar prophylls are also present at the bases
of the secondary branchlets (fig. 6c). Each of the secondary branchlets
apparently terminates in a solitary flower, with a bract and a similar bracteole
and no sign of any vestigial tip, but in some instances there was evidence of the
axis extending above the apparently terminal flower of the primary branches.
In fig. 6 a vestigial tip is shown, and in fig. 7 the apex of a branch is shown in
which the tip has developed and terminates in a flower, the penultimate flower
being sessile. This development of the tip is not common and was only observed
in one or two of the many branches examined.
The outer bract subtending each flower is, as described by L. Bolus, herbaceous
in the lower half, chartaceous round the margin, o-8-i cm. long (pi. XI, Sa).
The inner sterile bract or bracteole is similar but broader and rounded towards
the apex (fig. 8b). Between these bracts and the flowers what appeared to be
the membranous tips of a second pair of bracts could be seen in almost every
case. Fig. 8 is the terminal bud of the primary branch sketched in fig. 6, slightly
enlarged, showing the tips of the two inner bracts between the bud and the two
outer bracts. When the outer bracts were removed with care the traces of two
extremely delicate inner bracts were revealed. The position of these bracts is
shown diagrammatically in fig. 9. Within the outer bract, i, is a vestigial bract,
ii, which, like the outer one, partially envelops the second outer bract, iii.
Between the bract iii and the ovary is the trace of a fourth bract, iv. The two
vestigial bracts are shown in figs. Sc and Sd.
The inner of the two vestigial bracts was so delicate that it tended to
disintegrate when touched, but a number of them were examined and it was
found that at the base they were more or less adnate to the base of the ovary.
The ovary, with the remains of a bract attached, is shown in fig. Se. This
second bract is larger than the lower vestigial bract and might be compared
with the bracteoles of some species of Nivenia which are conspicuously longer and
wider than the subtending bracts, and are brown and membranous whereas the
bracts are firm and green.
These very delicate structures which are presumed to be traces of bracts are,
like those found in the Watsonia hybrid, pale pink and translucent, and when
moistened they immediately dissolved into a colourless mucilage. There is no
sign of any actual vestigial bracts in herbarium specimens which have been
examined, but in many of them the buds and exterior surface of the perianth
tube and lobes of open flowers have a shining, lacquered appearance. This
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 5 1
undoubtedly comes from the mucilage from the moistened vestigial bracts,
either due to rain having fallen on the inflorescence before the buds opened, or
to the process of drying and poisoning. The flowers of some herbarium specimens
of Watsonia pyramidata have the same lacquered appearance.
Assuming that these structures observed in Pillansia Templemanni are indeed
the vestigial traces of bracts, then the inflorescence in this plant fairly obviously
represents an intermediate stage between the spicate inflorescence of the
Ixioideae and a compound inflorescence such as that of Nivenia binata or N.
corymbosa. On pi. XI, in figs. 10 and n, are shown diagrammatically two
theoretical earlier stages in the development of the seemingly apical bud of the
primary branch of Pillansia Templemanni. If, in addition, the prophylls at the
bases of the primary and secondary branchlets, instead of being united and
sterile, were free and one or both of them subtended axillary branchlets, then
the inflorescence would scarcely differ from that of Nivenia corymbosa (Ker)
Baker. Weimarck, in his revision of Nivenia, illustrated the theoretical develop-
ment of the upper, umbel-like part of the main inflorescences of N. corymbosa
and N. binata Klatt. A plan showing the arrangement of the bracts and branches
at the first node of the main axis of an inflorescence of JV*. corymbosa which was
examined is given in fig. 12 on pi. XI. Bi is the first bract of the main axis
(marked X) and it subtends the lowest primary branch, 1 . The lowest internodes
of this branch are contracted and the two small bracts or prophylls at the base,
Bii and Biii, are thus extremely close together. The upper of these two bracts
is sterile but the lower one, Bii, subtends a secondary branchlet which, like the
primary branch, has its lowest internode contracted. It bears a tertiary branch-
let, 3, in the axil of the bract Biv. This in turn has a bract, Bv, at its base,
with a bud in its axil. Above the very short internodes at the bases of the
primary, secondary and tertiary branchlets are long internodes, but there is no
reason to describe these branchlets as this has already been done by
Weimarck.
The point that I wanted to illustrate in the diagram is the proximity of the
two bracts or prophylls at the base of the primary branch. Only a very slight
further abbreviation of the basal internodes would, in all probability, result in
the fusion of the overlapping margins next to the axis of the bracts Bii and Biii,
and the suppression of the bud in the axil of the lower bract (Bii) . This would
leave a primary branch with a pair of united sterile bracts or prophylls at the
base, between the branch and the axis, such as occurs in Pillansia Templemanni.
On the other hand, only one further stage in reduction in the branch of
Pillansia Templemanni shown on pi. XI, fig. 6, would result in the suppression of
the internodes of the secondary branchlets. If, instead of branchlets, flower-
buds were produced at these nodes, this would give a short lateral spikelet with
two sessile flowers each with its subtending bract and a pair of united prophylls,
or bracteoles, as they would now be called. The vestigial trace of a bract within
the bracteoles of one of the buds in the Watsonia hybrid (pi. XI, 36) seems to
suggest that a similar reduction has taken place in that plant.
52 ANNALS OF THE SOUTH AFRICAN MUSEUM
As observed by Haeckel, the apex of the branch generally disappears in an
early stage, but it occasionally happens that the uppermost flower is developed,
in which case, as recorded by Boehni, the flower is regular even if all those below
it are zygomorphic. I have observed this myself in a few specimens of Watsonia
Meriana.
6. The inflorescences of some species of Lapeyrousia
Another genus of interest from the phylogenetic and morphological view-
points, which requires a great deal more study, is Lapeyrousia. It is a fairly large
genus, well represented in South and tropical Africa, and in some species the
inflorescences present features which are associated with those found in the
Iridoideae and Crocoideae, while in others, probably the majority, the inflorescence
is reduced to the spicate type which is typical of the Ixioideae. In the subgenus
Sophronia, for instance, all the internodes of the main axis and the branches are
abbreviated and the leaves and flowers are congested in a dense 'sessile rosette',
as in Galaxia. In some species in the subgenus Ovieda intermediate stages between
a prolific, laxly branched inflorescence and a simple or branched spicate one
occur.
The subgenus Anomatheca is the only one of the three subgenera in which the
basal leaves arise from the corm, as is almost invariably the case in the Ixioideae.
The corms in this subgenus also differ in other respects from those of the other
two subgenera, in which the lowest green leaves do not arise from the corm
itself but from the aerial axis, usually shortly above the corm, as they do in
Galaxia and also all those members of the Iridoideae which have corms.
There is another feature of interest which is probably more conspicuous in
some species of Lapeyrousia than in any of the other genera. This is the matter
of adnation, which requires further investigation and has been mentioned before
in connection with the leaves. In several species of Lapeyrousia, notably L.
corymbosa, L. Fabricii and L. Jacquinii, it can be seen quite clearly that the branches
(also the flowers) with their subtending bracts are more or less adnate to the
axis. In L. Fabricii, a species which grows in sandy ground, the corm is deep-
seated and the portion of the aerial axis below the ground is frequently elongated.
Very often cormlets are produced in the axils of the lower leaves which arise
below the ground-level, one in each leaf axil. Such cormlets almost invariably
appear some distance below the node (see pi. XII, fig. i), breaking through the
leaf skin or lower portion of the sheath which is adnate to the axis.
In one or two specimens of L. Fabricii examined it was found that some of the
upper branches remain adnate to the axis for 6 or 7 mm. above the node, while
in one or two specimens of L. Jacquinii the lowest branch was found to emerge
from the axil of its subtending leaf below the node, rupturing the lower dorsal
part of the sheath as in the case of the cormlets mentioned above. In this species
also the rather large, conspicuously bicarinate and bifid prophylls are frequently
carried out of their normal position, at the base of the branch, and appear
half-way up the branch. The axis in these species mentioned above, as well as
Ann. S. Air. Mus. Vol. XL.
Plate XII
I, Lapeyrousia Fabricii (Thunb.) Ker, base of stem, with cormlets; i, node. 2, L. Jacquinii N. E,
Br., diagram of spike; i, bifid brae teoles. 3, L. Vaupeliana Dinter, diagram of top of inflorescence.
4, L. Bainesii Bak., diagram of top of inflorescence ; i, bifid bracteoles. 5, L. corymbosa (L.) Ker,
inflorescence, xf; i, prophylls; 5a, branchlet, X i|; 56, bract, Xi|; 5^, bracteole, xi|;
$d, bract subtending branch near top of inflorescence, X 6^. 6, L. corymbosa, branch of inflo-
rescence, the lowest flower with united bracteoles. 7, L. micrantha Bak., bract of branch near
top of inflorescence, X 6 J.
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
in several others, is often extremely flexuose, a condition which usually seems to
be associated with adnation. In some weak or drought forms of L. Jacquinii, in
which the lower branches are not developed, the axis is almost straight, but in
more robust plants with several branches it is extremely flexuose, the branching
often appearing divaricate as the primary branches are usually as long as the
main axis.
A diagram of the spike of L. Jacquinii is given on pi. XII, fig. 2, to illustrate
the adherence of the lower part of each bract to the axis, suggesting a sympodial
rather than a spicate arrangement. A vestigial tip is present and the bracteoles
are all distinctly bicarinate and bifid (forked in diagram). The fact that the
bracteoles of the apical flower here are also strongly bicarinate and bifid refutes
the argument of those botanists who maintain that this is due to compression
between the bud and axis.
L. Vaupeliana Dinter, from South West Africa, is among the tallest and is
undoubtedly the most richly ramified species in the genus. Some specimens in
the South African Museum Herbarium collected by Dinter [Dinter 2787 and
3374) are from 2 to 4 ft. high, and from shortly above the base upwards the axis,
which is distinctly flexuose, bears numerous branches, each in turn bearing
several branchlets, so that the whole plant is a dense mass of innumerable
branchiets, each as a rule terminating in a single flower with a membranous
bract and an identical bracteole. In one or two forms which are less vigorous
the upper parts of some of the lower branches are present only as abortive buds
and towards the top of the inflorescence on some of the branchlets there is a
penultimate sessile flower (compare the lateral branch in the diagram of L.
Bainesii on pi. XII, fig. 4). The upper part of an inflorescence of L. Vaupeliana
is shown diagrammatically on pi. XII, fig. 3. All the flowers in the upper part
of the inflorescence open together, those of the lower branches being still in bud.
The lower internodes of the primary branches are all elongated, so that in effect
the branching is more or less corymbose, but towards the ends of the branchlets
the internodes are almost equal and here the branching is dichotomous (see
diagram). In the lower part of the inflorescence there is a pair of united
prophylls at the base of the primary and most of the lower secondary branches,
but in the upper part these have not developed, although abortive buds can be
seen quite distinctly between the main axis and primary branches as well as
between the primary and lower secondary branchlets. With a lens it can be
seen in some cases that these buds have two collateral growing points.
The primary branches arise alternately from the axis, in accordance with the
distichous arrangement of the leaves, but the first secondary branchlet which is
developed is always in the axil of a bract which is on the same side as the
preceding bract or leaf. The same is true of the tertiary branchlets and finally
of the flowers. This arrangement is shown in the diagram, although in the
plants examined it is not immediately obvious owing to a slight spiral twist in
the stems. It can, however, be fairly readily traced as the stems are sharply
angled. This inflorescence might be compared with that of Nivenia corymbosa
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 55
with the following differences noted: (i) The prophylls at the bases of the
branches, when developed, are united and always sterile; (2) each flower has
a bracteole but there is no upper sterile bract.
In his very brief description of the species, which he stated was near L. Bainesii,
Dinter merely referred to the inflorescence as prolific and laxly branched. He
made no reference to the style branches which in all the specimens I have seen
are simple and not bifid as they are almost invariably throughout the genus.
The corm, described by Dinter as 'broadly bell-shaped', is typical of the sub-
genera Sophronia and Ovieda, flat at the base and broadly conical above, and the
lower leaves arise from the axis above the corm.
The inflorescence described above and the simple-style branches seem to
indicate that these characters are the most primitive in the genus. Comparing
the inflorescence of L. Bainesii with that of L. Vaupeliana, it is evident that the
former represents a transition stage between the much-branched inflorescence
of the latter and the spicate inflorescence which occurs in many species of
Lapeyrousia. A diagram of the top of an inflorescence of L. Bainesii is sketched on
pi. XII, fig. 4. In this the main axis terminates in a four-flowered spike. The
apical flower and the flower below it have single bracteoles, identical with the
bract, but the bracteoles of the two lower flowers are bifid (forked in the
diagram). The two branches have two flowers each, all of them with a single
bracteole. On the lower branch there is a slight indication of a vestigial tip
above the upper flower. The lower internode of both branches is long but the
upper one abbreviated, so that the two flowers are comparatively close together,
a circumstance which was commented on by Weimarck in connection with
L. plagiostoma.
This arrangement of the flowers in pairs is even more striking in L. corymbosa
Ker, a species fairly common on the Cape Peninsula and in some of the
neighbouring districts. In this species the upper internode of the branches is
almost completely suppressed and the two flowers are very close together (see
figs. 5 and 5a on pi. XII), resembling the arrangement which Weimarck
referred to as 'binate flowers'. In living material examined there was no visible
sign of the apex extending above the upper flowers and in practically every case
the bracteole was found to be a single member, slightly narrower than the bract
but otherwise identical, as can be seen in figs. 56 and 5c. In only one specimen
was a bifid bracteole found on one of the lower branches, where a reduction
had taken place and instead of a two-flowered branchlet there was a single
sessile flower with a pair of united bracteoles (see fig. 6) . On the lowest branch
of the plant figured in the diagram (XII, 5) the united prophylls have been
displaced and are carried out of the position they normally occupy, a circum-
stance referred to above in connection with L. Jacquinii.
The small, very much reduced bracts (about 1 mm. long) subtending some of
the upper branches of the inflorescences in L. corymbosa and L. micrantha, two
closely related, well-branched species, are shown in figs. $d and 7 on pi. XII.
The first shows a tendency towards tricuspidation and the second is distinctly
56 ANNALS OF THE SOUTH AFRICAN MUSEUM
tricuspidate. In the lower fully developed bracts in the same inflorescences there
is no sign of this lobing, which is probably due to reduction.
7. Reduction of inter nodes and the rhipidia in the more primitive genera
The most simple and primitive combination of characters in the South
African genera is found in the genus Aristea. The systematic position of this
genus and the confusion which ensued from its having been combined with the
genus Nivenia was discussed by Weimarck in his monograph. He placed it in its
correct position, in the tribe Sisyrinchieae, with Sisyrinchium and several other
genera, none of which, with the exception of Bobartia, are South African.
Weimarck regarded the binate rhipidia as the typical inflorescence type in
Aristea and the various deviations in the types of inflorescence he attributed as
a rule to reduction of flowers, so that bracts and prophylls remain sterile. My
own investigations seem to show that bracts and prophylls which have become
sterile through reduction either disappear or remain as sterile members which
do not again become fertile except in the rare case of proliferation such as has
occurred in Watsonia bulbifera. In Lapeyrousia Vaupeliajia, for instance, in spite of
all the ramification, the prophylls and bracteoles are all sterile, as they are
throughout the genus.
Probably the most primitive inflorescence to be found in the South African
Iridaceae is that which occurs in some of the larger species of Aristea, such as
A. Bakeri and A. capitata. In these two tall, robust species, with their compound
racemose branching, sterile prophylls are present only in some of the weaker
forms in which the branchlets have not developed. The branching of a fairly
well developed specimen of A. capitata is shown diagrammatically on pi. XIII,
fig. 1. The branches and branchlets throughout end in 'binate rhipidia', as
Weimarck has named this type of inflorescence. The lowest primary branch
bears four secondary branchlets and these sometimes give rise to tertiary branch-
lets, as is shown in the diagram. The lowest secondary branchlet arises at the
extreme base of the primary branch, next to the axis. The lateral branches
become progressively shorter towards the top of the axis, but as can be seen in
the third, fourth and fifth, each still bears two or three branchlets, although in
these the two lower internodes are extremely abbreviated and the two lower
branchlets arise next to the axis. In the diagram it is only possible to show one
plane, but the actual position of the branchlets on either side of the branch is
shown in fig. \a. This reduction of the internodes and position of the bracts or
prophylls is significant, for only a very slight further contraction of the basal
internodes, or the adnation of these abbreviated internodes to the axis, would
almost undoubtedly result in the fusion of the bracts and the suppression of one
or both of the buds.
The end of a branch of a recently described species of Aristea (A. latifolia
Lewis), slightly enlarged and without the flowers which terminate each branchlet
or pedicel, is shown on pi. XIII, fig. 2. This tall and well-branched species,
somewhat similar in appearance to A. Bakeri but quite distinct in many respects,
Ann. S. Afr. Mus. Vol. XL.
Plate XIII
i, Aristea capitata Ker, diagram of inflorescence; \a, third node of inflorescence, bract removed.
2, Aristea latifolia Lewis, binate rhipidia, x 2 (the bract and two bracteoles removed from the
rhipidium on the left) ; 2a, bracteole, X 2 ; 2b, plan of binate four-flowered rhipidia. 3, Aristea
oligocephala Baker, binate rhipidia with vestigial tip, x f; 30, same, with one rhipidium removed;
36, bracteole.
58 ANNALS OF THE SOUTH AFRICAN MUSEUM
grows only at fairly high altitudes (about 4,000-5,500 ft.) on mountains in the
Paarl, Worcester and Caledon Divisions. It was selected for illustrating the
inflorescence as the bracts were small and narrow and the internodes slightly
longer than in any of the other species. In the figure the outer bract and two
succeeding bracteoles on the left are removed and only the upper one, which is
sterile in this case, is shown. Except that it is very slightly smaller, there is no
difference at all between the first bracteole, shown in fig. 2 a, and the bract.
There is a slight indication of an apex between the two rhipidia.
The inflorescences of several species of Aristea were examined and as a rule
there is little if any visible sign of an apex between the two rhipidia, but in some
specimens of A. oligocephala an actual vestigial tip was found. This is illustrated
on pi. XIII, fig. 3. Fig. 3A is the same, slightly enlarged, with one rhipidium
removed. This vestigial tip was only found in three inflorescences out of about
fifty that were examined ; in one of them it was more robust and terminated in
a normal flower. One of the strongly carinate bracteoles with its wide mem-
branous, colourless margins is shown in fig. 3^.
A point that should be mentioned in connection with the rhipidia of A. latifolia
is that the flowers are more or less spirally arranged. A plan showing the arrange-
ment of a pair of four-flowered rhipidia in this species is given on pi. XIII, 20.
In each rhipidium the first and second flowers are more or less opposite and the
third and fourth opposite each other and almost at right angles to the first pair.
The fifth bracteole in each is sterile. The resemblance of these rhipidia (if this
term is indeed the correct one in this case) to the umbellate inflorescences of
the Amaryllidaceae, with their condensed bostrychoid cymes, led to a compara-
tive examination of some of these umbels. The first one available was Tulbaghia
violacea Harv., a species from the eastern Cape Province, and as this happened
to provide some further evidence of the fusion of the bracteoles it will be
described briefly here.
The genus Tulbaghia is one which Hutchinson, in his Families of Flowering
Plants, transferred from the family Liliaceae to Amaryllidaceae on account of
its umbellate inflorescence. In nearly all the inflorescences of Tulbaghia violacea
examined the umbel consisted of two cymes, usually with four or five perfect
flowers in each and one or two abortive buds. In one specimen there were three
such cymes and a trace of a fourth. In all of them the inner bracts, or the
bracteoles, were variously united, with only the short acute tips free, either one
free and three or four united, or in pairs, or all united. In no case were they all
free. The first specimen examined was slightly abnormal, with the pedicels of
two flowers in each of the two cymes united up to the base of the perianth. This
specimen is illustrated on pi. XIV, figs. 1 and \a. Fig. \b shows four normal
buds with their united bracteoles, and in fig. ic these same bracteoles are shown
opened out, with the buds removed. A diagram of the arrangement of the
flowers and bracteoles in one of the umbels is shown in fig. id.
It has been observed by Arber and others that the conditions of pressure under
which lateral buds arise in some of the Monocotyledons sometimes results in
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE
59
anomalous fusions. Such a fusion has occurred in this species of Tulbaghia. In
the umbels of other members of the Amaryllidaceae examined, such as Crinum,
Cyrtanthus and JVerine, the bracteoles are all free but they are very much reduced
in size and are all more or less filiform and functionless, the function of protecting
the buds being performed by the much enlarged outer bracts or spathes.
In Bobartia, the other South African member of the tribe Sisyrinchieae, the
discovery during the past twelve years of two very distinct species is of particular
interest, as they provide some important 'missing links'. These species, B.
paniculata Lewis (Journ. S.A. Bot. VII, 50 (1941)) and B. lilacina Lewis {Journ.
S.A. Bot. XI, 108 (1945)), both from mountain districts in the Cape Province,
u
1a
Plate xiv
Tulbaghia violacea Harv. ; 1, umbel, X-f; i, united pedicels; la, two flowers with united pedicels,
xf; 1 b, four united bracteoles subtending four flower-buds, X
same bracteoles with buds
removed, xf; id, plan of umbel with bracteoles variously united.
60 ANNALS OF THE SOUTH AFRICAN MUSEUM
will necessitate some alteration to the definition of the genus of which the
inflorescence has hitherto been described as compact or unbranched. It might
be mentioned here that during a visit to Kew and some herbaria in Europe in
1949 it was found that the laxly branched species named B. Marina was actually
first collected by Drege over a hundred years ago. The extremely delicate and
fugacious pale mauve flowers are missing from the specimens {Drege 1538) in
the herbarium at Kew and in Paris. Drege and Meyer placed this plant in the
genus Moraea, but it was not described or named.
Bobartid lilacina is a laxly branched species with slender stems and small bracts
and prophylls, the latter united, distinctly bifid and bicarinate. In most cases
the prophylls are sterile but occasionally a branchlet develops in the axil of the
upper of the two united prophylls. Two such branchlets are shown in a diagram
of the inflorescence of B. lilacina in fig. 3 on pi. XV. In B. paniculata the one
to three slender branches are borne near the top of the stem. It is evident that
in this species a considerable reduction has taken place in most of the internodes.
There is little doubt that B. lilacina is the most primitive species in the genus and
that the inflorescence of B. paniculata represents an intermediate stage between
the laxly branched inflorescence of B. lilacina and the congested type which
occurs in the other twelve to fifteen species, in which a further reduction has
taken place in the upper internodes of the main axis and the lateral
branches.
On pi. XV, fig. 1, is illustrated an inflorescence (without flowers) of B. gladiata
Ker, and in fig. 1 a a diagram of the same inflorescence. This is more or less the
typical congested type of inflorescence most common to the genus ; in some
species it is very much denser and in others, such as B.filiformis, more reduced,
but the arrangement is more or less the same throughout, except in B. paniculata
and B. lilacina. The internodes are all extremely abbreviated and the lowest
bract, which is sometimes much enlarged, often appears as a terminal extension
of the axis, the inflorescence above this bract being forced out of position and
more or less at right angles to the lower part of the axis. As can be seen in the
diagram, however, the apparently terminal bract is actually lateral, with a bud
in its axil which, in more robust forms of B. gladiata, frequently develops. In
B. indica L. the lateral position of this bract is more evident and the considerably
denser inflorescence occupies the terminal position.
The thickened lines in fig. 1 represent the strongly bicarinate prophylls, which
are forked in the diagram. Fig. 2 is the same as fig. \a but with the internodes
extended and with the two lower buds developed into branches which are
indicated by dotted lines, as well as a branchlet in the axil of one of the pairs of
prophylls. With the addition of these branches it represents a diagram of the
inflorescence of B. paniculata.
Through B. lilacina the genus Bobartia is more closely allied to Aristea. It differs
from Aristea, among other things, in having the prophylls united throughout,
even in the rhipidia. In this respect the rhipidia of Bobartia are the same as those
of Homer ia, Ferraria, Moraea, Dietes, etc., and as some of them were examined in
Ann. S. Afr. Mus. Vol. XL.
Plate XV
Bobartia gladiata (L. f.) Ker, inflorescence; la, diagram of same inflorescence. 2, same as la
>ut with internodes extended; with addition of branches marked with dotted lines this represents
the inflorescence of B. paniculata Lewis. 3, B. lilacina Lewis, diagram of inflorescence.
62 ANNALS OF THE SOUTH AFRICAN MUSEUM
some detail this occurrence of the bicarinate bifid prophylls or bracteoles should
be discussed.
In all these genera (i.e. Bohartia, Moraea, etc.) the lowest bract or spathe of
the rhipidium completely encircles the inflorescence or rhipidium, and has a
short, closed sheath at the base. The second bract is almost always longer and
broader and also encircles the inflorescence, but the margins, although they
overlap at the base, remain free. In none of the specimens examined was this
bract found to be bifid or bicarinate and there is little doubt that it is a single
bract. Each of these lower bracts subtends a flower, but the upper flowers in
the rhipidia are subtended by more or less bicarinate and often distinctly bifid
bracteoles.
There seems every reason to believe that, just as the prophylls of the branches
in Bobartia, Moraea, Homeria, etc., consist of two united bracts, the lower of which
is sterile (often both are sterile), the same is true of the bracteoles in the rhipidia
of these genera. That such fusions of two overlapping members can and do
take place, due to the reduction of internodes and conditions of congestion and
pressure, has already been shown. In the plan of the Aristea rhipidia on pi. XIII,
fig. 2b, the somewhat spiral arrangement of the flowers and overlapping of their
bracteoles is illustrated, and reference was made to the fact that the internodes
in this species are slightly longer than is usual in the genus. A contraction of the
internodes in a similar spirally arranged inflorescence might well have resulted
in the fusion of the bracteoles and consequent suppression of the lower of the
two buds, and it is possible that such a fusion having once taken place it has
become an hereditary character.
I am also of the opinion that the rhipidia in these genera (Bobartia, etc.) are
derived from binate rhipidia through a closer adnation of the two lateral
branches, resulting in the complete disappearance of the apex and of all except
the first flower of the lower of the two rhipidia, the flowering being continued
only in the upper one. This seems to be the only explanation for the structure of
the rhipidia, in which each of the two lower flowers is subtended by an outer
bract, or spathe, whereas the upper flowers are all subtended by united
bracteoles.
A diagram illustrating this theory is given on pi. XVI. Fig. i on pi. XVI is a
diagram of the inflorescence of Dietes bicolor (Lindl.) Sweet. The first flower is
subtended by the lowest bract, the second by the second much larger and longer
bract, and those above by bifid bicarinate bracteoles (forked in the diagram).
Fig. i a is a plan of the same inflorescence showing the relative positions of the
bracts and flowers. (The arrangement here is exactly the same as in Iris
pseudacorus as shown by Haeckel.) Fig. ib is a theoretical diagram of the same
inflorescence but with the internodes slightly extended and with a vestigial tip
added in the position I presume that it would occupy. For comparison I have
added a diagram (fig. 2) of the actual inflorescence of the Aristea shown on
pi. XIII, fig. 2. It has already been pointed out that in Aristea there are only
traces of a vestigial tip between the two rhipidia in a very few cases. In most of
tf
Ann. S. Afr. Mus. Vol. XL.
Plate XVI
i, Dietes bicolor (Lindl.) Sweet, diagram of rhipidium: la, plan of same inflorescence; ib,
theoretical diagram of same inflorescence with internodes extended and a vestigial tip (i) added.
2, Aristea latifolia Lewis, diagram of binate rhipidia shown in fig. 2 on pi. XII. 3, Moraea ramo-
sissima (L. f.) Druce, part of inflorescence; ^a, diagram of branching at the lower node in fig. 3,
showing bracts and short and long internodes; 36, diagram of branching from the same node
but with the lower internodes of the branches extended; 3c, plan of bracts and branches at the
same node.
64 ANNALS OF THE SOUTH AFRICAN MUSEUM
the species in that genus there is no visible sign of a tip and the two rhipidia are
extremely close together.
It is stated above that in Bobartia, Moraea, etc., the lower of the two fused
prophylls of the branches is always sterile, but an exception was found in an
unusually tall and robust specimen of Moraea ramosissima; in this branchlets
were present in the axils of both of the lowest prophylls.
Moraea ramosissima (L.f.) Druce is usually 2 to 4 ft. high, and the very apt
specific epithet refers to the numerous branches and branchlets. A part of the
southern area of the Gape Peninsula which had been completely burnt out a
few months previously was visited in 1950 and in a small marshy area there was
a dense growth of this species, many of the plants 6 to 7 ft. high. On examining
one of these plants two branchlets were found in the axils of two partly united
prophylls. A sketch of the upper part of the axis with the branching at two of
the nodes is given on pi. XVI, fig. 3. Fig. 30 is a diagram of the branching at
the lower of these two nodes, which illustrates the congestion of the branchlets
and their subtending bracts and the succession of short and long internodes.
The lowest bract, B, subtends the primary branch, 1. Bracts Bi and Bii,
although shown as separate members, are actually partly united and each
subtends a secondary branchlet (numbered 2 and 3). Biii represents a pair of
united bracts subtending a single tertiary branchlet, 4, and Biv also united
bracts with a single branchlet, 5. Between the fourth and fifth branchlets is a
bud. Each branchlet terminates in a two- or three-flowered rhipidium.
Fig. 3b represents the branching from the same node but with the lower inter-
nodes of the branches extended to show the arrangement more clearly. Since
only one plane can be shown in these diagrams I have added a plan (fig. 3) to
show the position of the branches and bracts in relation to the axis (marked x) .
The base of the bract subtending the primary branch encircles and forms a
short closed sheath round the axis.
8. Conclusion and a reference to the value of the bracts as a diagnostic character in
taxonomic work
The various inflorescences which have been discussed in the preceding pages
all provide some evidence in support of HaeckePs assertion that the more
composite types of inflorescences in the Iridaceae are derived from a primitive
much-branched type. Reductions in the number of flowers and suppression of
some of the internodes have finally resulted in the simple or branched spicate
inflorescence of the Ixioideae. The fusion of the prophylls has been shown to be
due to the reductions of internodes, as well as in part to the width of the sheathing
bases of these bracts so that, when reductions have taken place, these broad
sheathing bases have first overlapped, then finally become united.
These conclusions are not in accordance with Weimarck's theory that the
spicate inflorescence of the Ixioideae is the most primitive arrangement in the
family, and that the more complex types of inflorescence were derived from this.
Two or three factors seem to preclude the correctness of this theory. One is the
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 65
presence of the bifid or bicarinate bracteoles in the spicate inflorescence of the
Ixioideae. This type of spicate inflorescence occurs only in the Ixioideae, which
is generally considered to be the most highly evolved of the three groups into
which the family has been divided (the development of the corm, leaves,
inflorescence and flowers all bear this out). Weimarck's assumption that the
type of spike in the Ixioideae was the most primitive and simple arrangement,
would seem to imply that those genera with more complex inflorescences, such
as Aristea, Nivenia, Bobartia, etc., which according to Weimarck are evolved from
this type of spike, were all at a more advanced evolutionary stage than those in
the Ixioideae, such as Gladiolus, Antholyza, etc.
There is little doubt that the reduction in the number of flowers in the rhipidia
or of some of the internodes, such as has occurred in some species of Aristea as
well as in other genera, is associated with other variations, such as either the
enlargement of the bracts and prophylls or, in some species in which the pedicel
is reduced and the flower almost sessile, this is accompanied by an elongation of
the ovary or, in those genera in which the pedicel has entirely disappeared, the
development of a perianth tube. Throughout the Ixioideae the flowers are sessile
and all have a perianth tube, even though in one or two of the genera, such as
Melasphaerula and Geissorrhiza, it is very short. The reduction in the number of
flowers has also resulted in the formation of flowers of very much firmer texture
with an anthesis of two or three days instead of only a few hours. These
apparently correlated variations will be referred to again when discussing the
flowers.
The bracts and bracteoles, although occasional exceptions do occur, on the
whole provide fairly reliable diagnostic characters for use in taxonomic work
and deserve more attention than they have so far received. In the majority of
the genera in the Ixioideae with a spicate inflorescence there is some evidence
(though sometimes barely perceptible) of the bracteoles being bifid or bicarinate
and in nearly every case they are either slightly or distinctly shorter than the
bracts. One group of closely allied genera has been mentioned in Part I in
which this order is reversed In Anapalina, Exohebea, Tanaosolen and Tritoniopsis
the bracteoles are nearly always longer than the bracts (in Anapalina triticea
and A. Burchellii considerably so) but otherwise they do not differ from them.
The membranous tricuspidate bracts of Ixia and Tritonia subgenus Dichone, and
of one or two other genera, have also been mentioned in Part I.
The vestigial traces of bracts in Pillansia Templemanni and the species of
Watsonia discussed and the occurrence of cormlets in Watsonia bidbifera seem to
indicate that these plants are of comparatively recent origin.
Another reduction probably of fairly recent origin is that of the terminal
spike in Antholyza ringens L. In this species the long terminal axis overtopping
the short lateral spike usually has near its apex only a single barren bract, but
occasionally it reverts to its ancestral form and there are two or three bracts,
sometimes even a small flower. This monotypic genus is so closely allied to the
genus Anaclanthe N. E. Br, that I doubt whether the latter should be maintained
66 ANNALS OF THE SOUTH AFRICAN MUSEUM
as a separate genus. In Anaclanthe spicata (Thunb.) N. E. Br. there are three or
four short lateral branches with secund spikes borne more or less at right angles
to the axis, which terminates in a spike, but apart from this there is no real
difference between the two genera of which the corms, leaves and bracts are
almost identical . The flowers of Anaclanthe spicata are slightly smaller than the
typical flowers of Antholyza ringens but the size of the plants and flowers of the
latter is extremely variable, and in the smaller forms there is little if any dif-
ference to be seen in the flowers of these two plants. The disappearance of the
terminal spike in Antholyza ringens is probably due to the comparatively recent
adaptation of these flowers to bird-pollination, as a result of which the flowers
at the top of the spike, which were not so easily accessible to visiting birds, tended
to remain unfertilized and finally became sterile and have now almost entirely
disappeared.
Part III. The Subterranean Stems
i. Introductory remarks
In his Classification of the Flowering Plants Rendle observed, regarding the corms
of the Iridaceae, that 'the corm is especially characteristic of the Iridaceae,
varying in size in different genera and species. The size of the corm, its shape
and more especially the character of the sheathing scales, afford means of
distinguishing the very numerous species of Crocus and Gladiolus.''
Except for the shrubby genera and three or four which have rhizomes, such
as Aristea, Dietes, etc., the great majority of the South African genera have
corms, and Rendle's observation is very apt, for those who are familiar with this
family can frequently recognize the genus, or in some cases the species, or
group of species, to which a plant belongs by the characteristic shape or distinct
fibrous tunics of the corm. This is not always the case, however, for since the
character of the fibrous tunics or sheathing scales of the corm is naturally
associated with those of the leaves and bracts, so it is only to be expected that
genera which have very similar or identical leaves and bracts should also have
very similar or almost identical corms. Examples of some genera which it is
difficult or impossible to distinguish by the corm alone are: Synnotia and
Sparaxis ; Lapeyrousia subgenus Anomatheca and Freesia ; Exohebea and Anapalina ;
Ixia and Tritonia subgenus Dichone; Babiana, Anaclanthe and Antholyza; Homo-
glossum and some species of Gladiolus. In all these cases the respective genera
linked together are very closely allied and are separated mainly on account of
differences in their floral characters.
Although there is little doubt that they furnish some distinct evidence of
phylogenetic trends in the family, up to the present comparatively little use has
been made of the character of the corm in taxonomic work, nor has the mor-
phology of these organs been studied in much detail. Foster, in his revision of
Geissorrhiza, has used the nature of the corm tunics, which he observed to be
associated with other morphological features, as a diagnostic characteristic.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 67
Hutchinson made some use of the nature of the underground stems in the keys
to the tribes and genera of the Iridaceae in his Families of Flowering Plants, and
to some extent I have also made use of the character of the corm, in combination
with other characters, in a revision of the Iridaceae of the Gape Peninsula.
One obvious drawback to relying too much on the character of the corm or
rhizome alone in taxonomic work is the fact that in herbarium material, more
especially in the older collections, these organs are not always present. In 1949,
when examining some of the old collections in England and Europe such as
those of Linne, Thunberg, Burmann, Lamarck and others, it was found that in
hardly any of the specimens are the corms or rhizomes present. Though this is
regrettable, as so many of them are type specimens, it is hardly surprising, since
Linne and the early botanists following him used only the floral or sexual
characters as a means of identifying the families and genera.
An examination of some of the subterranean stems of the South African
Iridaceae has shown that more than one type of corm occurs and that there are
one or two intermediate stages between a rhizome and corm. The swollen
underground stems of several species of the shrubby genera have been described
by Adamson and will not be described here, but some of the structures in
various other genera which have not hitherto been described in any detail
are mentioned in the following pages.
2. Transition from rhizome to corm
Of the forty to fifty genera of Iridaceae which occur in South Africa, the three
shrubby genera form a small, distinct and fairly homogeneous group, with
woody aerial stems with secondary thickening and swollen more or less sub-
terranean basal stems in which secondary thickening also takes place. Many of
the species in these genera grow in wet or fairly moist situations. Of the other
genera only four have been recorded as having rhizomes, the remainder, which
constitute the great majority of genera and species, having a corm or an under-
ground storage stem which approaches a corm in structure.
Three of the genera with rhizomes are in the Iridoideae, namely Aristea,
Bobartia and Dietes, and all these are evergreen herbs. Schizostylis, the fourth
genus with a rhizome, belongs to the Ixioideae and differs from the other three
genera in losing its leaves, together with the aerial axis, at the end of the flowering
season. The subterranean parts of the plant are probably the last to be affected
by the various changes which take place in the process of evolution and it is
very possible that in Schizostylis the habitat may account for the persistence of
the rhizome. The two closely allied species (perhaps only one species and a
variety) are found only in wet situations, along the margins of streams, where
the subterranean stem is not subject to any great seasonal fluctuations in
moisture. This genus differs from the fairly large and widely distributed genus
Hesperantha only in having a rhizome instead of a corm. The corms of Hesperantha,
which seem to provide some evidence of having been derived from a rhizome,
are discussed later.
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
Arber suggested that there was some degree of probability that in the Iridaceae
the rhizome may have preceded the bulb. So far no bulbs have been found in
any of the South African genera, but there seems little doubt that the more
primitive organ is the rhizome and that the corm is a later development, in all
probability evolved as a result of changes in climatic conditions. With the
exception of Schizostylis which is found only in wet places, the few South African
genera which have rhizomes are evergreen, and their distribution is significant
for they are found only in the coastal or mountain districts with a fairly high
and regular rainfall, and not in the dry regions such as the Karoo or Namaqua-
land where only the geophytic genera with corms, in which the aerial portions
die away at the end of each season, are able to survive.
Of the South African Iridaceae the most simple and primitive combination
of characters occurs in the genus Aristea, in which the subterranean stem in all
the species in which it has been seen is either a short, somewhat woody rhizome
with numerous, very short internodes, or else more slender and elongated, with
longer internodes and more in the nature of a sucker. The leaves in most of the
species are the simple isobilateral equitant type which Arber regards as being
the most primitive in the family. In Bobartia, the other South African member of
the Sisyrinchinae (in the Iridoideae) , most of the species have a somewhat woody
rhizome, but in some of them, instead of growing horizontally, the underground
stem is vertical and more swollen and in structure it shows distinct indications
of a transition to a corm. Bobartia Marina Lewis, which is considered to be the
most primitive member of the genus (see Part II), has a rhizome which scarcely
differs from that of a few species of Aristea. In Dietes, the other member of the
Iridoideae with a rhizome, the rhizome is also very similar to that of some species
of Aristea, i.e. rather woody with short internodes.
The type of corm which occurs in some genera in the Iridoideae will be dis-
cussed later, as it differs in some respects from the type which is most common
in the Ixioideae. In most of the genera in the latter group the abbreviated portion
of the aerial axis below the ground becomes swollen and forms a new corm each
season, while in almost every case the previous season's corm dies away as its
food reserves are used up, though it frequently remains at the base of the new
corm as a flat, hard, dry structure. There are three exceptions in this group,
namely Schizostylis, which has been mentioned above, Dierama, in which the
corms are perennial and the leaves evergreen, and Pillansia, which is also
evergreen.
The systematic position of the genus Pillansia has already been mentioned in
connection with the inflorescence. On account of certain features observed in
the inflorescence it is considered to be intermediate between the Iridoideae and
Ixioideae, and in the subterranean stem of this plant there is also evidence of a
link between these two groups, for it presents a rather striking transition from
a rhizome to the type of corm which is most common in the Ixioideae. Each
year a swollen corm-like structure is formed which perennates, and the single
long, green, isobilateral type of leaf to which it gives rise is persistent for two or
Ann. S. Afr. Mus. Vol. XL.
Plate XVII
1-4, Pillansia Templemanni (Bak.) L. Bol. 1, part of underground stem, xf; i, old flowering
axis; ii, base of dead leaf; iii, bases of green leaves; iv, base of flowering axis; v, sheathing
leaves. 2, longitudinal section of part of underground stem, with leaves removed, X f ; i, short
scale leaves. 3, part of underground stem and base of aerial stem, with outer leaves removed,
xf; i, base of leaf; ii, short scale leaves; iii, bud; iv, leaf scar; v, scar of sheathing leaf.
4, transverse section of root, x 16; 4a, t. s. of part of root, X50; i, cork; ii, epidermis ; iii,
cortex; iv, endodermis; v, pericycle; vi, phloem. 5, Dierama pendulum (L. f.) Baker, longitudinal
section of corm, X i-|; i, contractile roots.
70 ANNALS OF THE SOUTH AFRICAN MUSEUM
three years. The growth of the underground stem is in a horizontal direction
and in many ways it approximates more closely to a rhizome than a corm.
A portion of one of these subterranean stems is shown in fig. i on pi. XVII,
with the fibrous tunics removed. This represents only a fifth of the actual
specimen in which there were twenty-five swollen corm-like bodies, half of them
lying immediately below the portion illustrated. The short sheathing leaf
produced near the base of each 'corm' does not become detached but, after the
soft living tissues have died and broken down, the coarse fibrous strands continue
to form a protective sheath around the corm. The second leaf produced is the
long green one and above this there are two or three very short brown scale
leaves, similar to those usually found on rhizomes. The position of these small
scale leaves is shown in figs. 2 and 3, the former a longitudinal section of two
of the corms and the latter one of the corms and the base of the flowering axis,
with all except the scale leaves removed. (For convenience these two are shown
in a vertical position but they should actually be horizontal.) The portion of
the axis which gives rise to the scale leaves is so very much reduced that no
internodes can be discerned in this region.
Each corm-body has a single, rather large root with a polyarch cylinder (see
fig. 4) and very large vessels. A transverse section of a portion of the root,
showing the anatomy in more detail, is given in fig. \a. The cells of the cortex
are pitted and of the endodermis much thickened, and the outer walls of the
epidermis are suberised. The vascular bundles of the well-defined stele of the
underground axis are concentric.
A section of a corm of Dierama pendulum, * the other evergreen member of the
Ixioideae, is shown for comparison on the same plate (fig. 5). As in the case of
Pillansia this also was probably derived from a rhizome but it represents a later
transition stage and differs in several respects from the simpler corm-like
structure of Pillansia. The corm illustrated was attached to nine living corms
growing more or less in a vertical direction one above the other and forming a
dense mass. In order to maintain its position or level in the ground the few
adventitious roots developed at the base are strong and contractile. The corm
gives rise to several leaves, all of which, even the outer ones, are the simple
linear isobilateral equitant type with no secondary winging or pseudo-mid-rib,
and there are no distinct scale or sheathing leaves. They differ in behaviour,
however, for the inner leaves, which are more swollen at the base than the outer
ones, become detached at the base, possibly through the formation of an absciss
layer, although they remain held in position by the outer leaves which remain
attached to the axis after they have died. These outer leaves, although similar
in appearance to the inner ones, are probably more in the nature of elongated
sheathing leaves.
The corms in the majority of genera in the Ixioideae are of the same type as
that of D. pendulum. They differ only in being annual instead of perennial and
in having the outer leaves reduced to short leaf-sheaths. In all of these the corm
* According to N. E. Brown the name Dierama is neuter, not feminine.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 7 1
is formed from the abbreviated base of the aerial axis, with two or three or more
swollen internodes and a well-defined stele, the reproduction of the plant being-
maintained by the development of the bud in the axil of the upper leaf at the
commencement of the next growing-period.
As far as is known, all except one of the species of Dierama are evergreen. The
exception is D. trichorhizum (Baker) N. E. Brown, in which the previous season's
corm dies away as the new one develops. Through this species the section
Morphixia of the genus Ixia is closely connected with Dierama and in fact D.
trichorhizum was originally described as an Ixia by Baker. It is of interest to note
that although so closely related these two genera have quite different distribu-
tions which scarcely overlap. Dierama covers a wide area extending from
Tropical Africa to the Transvaal, Natal and eastern Cape Province and as far
south-west as the Humansdorp Division, while Ixia is confined to the south-
western districts of the Cape Province, with one or two species in Namaqualand
and the Karoo and only one extending eastwards as far as Port Elizabeth.
The transition from a perennial corm, such as that of Dierama, to an annual
one is probably due mostly to changes in climatic conditions, but it may also
to some extent be brought about by the conditions of extreme congestion which
result from the persistence of the older corms. These are pushed out of position
by the growth of the new corms and although they continue to live for several
years their lower buds appear to remain dormant and they do not produce new
leaves or flowering shoots. These congested conditions may have resulted
eventually in the formation of smaller corms, often with fewer leaves, followed
by the complete absorption of the food reserves of the older corms as the new
ones developed and the dying-off of the leaves and older sterile corms. In the
Pillansia specimen described, although twenty-five corm-like structures were
present, more than half of the older ones were doubled up beneath the newer
ones above and although these structures still contained living tissue their buds
remained dormant and no longer produced leaves or made any further growth.
In this respect they differ from the perennial swollen subterranean basal stems
of the three shrubby genera, in which buds continue to develop and give rise to
new aerial shoots for an undetermined number of years.
Two views of one of the annual types of corm are shown in figs. 6 and 6a on
pi. XX. This is a corm of Chasmanthe floribunda and is among the largest of the
corms of this type found in the Ixioideae. It is depressed-globose and bears
several basal leaves with fibrous bases which are persistent on the corm during
the dormant period (the 'tunics'), though the upper green parts of these leaves
die off and disappear at the end of the flowering season. Fig. 6 is a view of the
top of the corm, with the fibrous tunics removed ; the dark rings are the scars
left by the bases of the tunics. Fig. 6a is a longitudinal section of the corm,
showing the large, well-defined stele and the bud at the top, on the left of the
scar of the old flowering axis, which will give rise to the new flowering shoot at
the beginning of the next season. In this species one or two of the outer or lower
buds sometimes develop at the same time and give rise to new plants. As in the
72 ANNALS OF THE SOUTH AFRICAN MUSEUM
case of Dierama pendulum, the adventitious roots produced at the base of the corm
at the beginning of the growing period are strong and contractile.
3. The tuberous 'corm! ofFerraria
The very distinct subterranean stem of Ferraria, which probably represents a
transition stage from a rhizome to the type of corm which occurs in many genera
in the Iridoideae, such as Hexaglottis, Moraea, etc., differs from all others in the
family in having no persistent scale leaves or tunics, and is actually more in the
nature of a tuber. The underground parts of two plants of Ferraria ferrariola
(Jacq.) Willd. are shown in figs. 1 and \a on pi. XVIII. In this species the
growth is in a horizontal direction, a new, more or less rounded tuber being
formed each season while the older tubers of the two or three preceding years
to which it remains attached become somewhat flattened as they begin to
shrivel. The bud which develops at the beginning of each season is at first
enclosed in a short sheathing leaf, but this becomes ruptured as the internode
immediately above it begins to swell and usually in the older tubers no trace of
it remains.
More material of another species, F. undulata, was available for a more detailed
examination, and it was found that the tuber in this species is perennial, and
for three or four succeeding years a new bud develops on the upper side of the
slightly compressed body. A young tuber with its first shoot is shown in fig. 2
on pi. XVIII. Each shoot has three colourless and slightly fleshy sheathing
leaves, produced at the base of the bud, and above them is a fourth, longer,
and half-green sheathing leaf which is followed by one or two long isobilateral
equitant green leaves. A new tuber is formed by the swelling of the internode
between the three basal sheaths and the next two or three leaves. A longitudinal
section of a young tuber and shoot showing the position of the leaves and the
internode which develops into the new tuber is illustrated in fig. 2a.
The basal sheaths consist of parenchymatous tissue, without chlorophyll, and
small vascular strands without any sclerenchyma, and as the tuber increases in
size it bursts through these sheaths, which shrivel and disappear. Traces of
them can be seen at the base of the tuber in fig. 2a. A few ordinary adventitious
roots develop immediately above the basal sheaths of the young shoot (shown
in a transverse section of a bud in fig. 3) and a single, larger adventitious root,
which is contractile, slightly higher up. A contractile root starting to develop
is shown in the transverse section in fig. 3a; part of it can also be seen in fig. 3.
The leaves and roots borne by the shoot die off and disappear at the end of the
first season and no further leaves or roots are produced on the tuber itself, which
is probably able to absorb water.
The succeeding buds which develop on the tuber are not adventitious but
arise in the axils of the original leaves, of which only the scars remain, as can
be seen in fig. 4. The bud shown towards the base in this tuber is apparently
not in a leaf axil and might be adventitious but it is more likely that it was
originally in the axil of the upper of the three basal sheaths and was carried up
Ann. S. Afr. Mus. Vol. XL.
Plate XVI IT
i, Ferraria ferrariola (Jacq.) Willd., new corm attached to old corm, xf; la, new corm and
three old corms. 2-6, Ferraria undulata L. 2, young corm, X \\\ i, colourless sheaths; ii, half-
green sheath; iii, leaf; iv, contractile root; v, ordinary adventitious roots; 2a, longitudinal
section of young corm, X 3^; i, contractile root; ii, outer sheathing leaves; iii, leaf; iv, new
corm; v, ordinary adventitious root; vi, remains of old sheaths. 3, transverse section near base
of new corm, X7; 3a, t. s. near top of same, X7; i, contractile root; ii, adventitious roots;
iii, colourless sheathing leaves; iv, base of leaf. 4, older corm, from above, X2; i, scars of
shoots; ii, bud. 5, corm with shoot attached to older corm,
adventitious roots. 6, mature corm with flowering shoot, X f .
i, contractile root; ii, ordinary
74 ANNALS OF THE SOUTH AFRICAN MUSEUM
out of position by the swelling of the tuber. A new tuber is formed by each bud,
which either remains attached to its parent, as shown in fig. 5, or else the thin
connection between the two tubers is severed by the action of the contractile
root of the new shoot. Sometimes two or three tubers develop in this way,
somewhat obliquely one above the other, and in this case no further develop-
ment seems to take place in the lower tubers, only the upper producing a new
shoot or flowering axis. The tubers in this species grow in a more or less vertical
position but are frequently pulled into a horizontal position by the contractile
root.
It is only after the third or fourth year that the tuber produces an inflorescence,
after which it begins to shrivel (see fig. 6). A few ordinary adventitious roots
are produced at the base of the flowering shoot, but no contractile roots, and
the shoot remains attached to the old tuber until it withers and finally disappears
at the end of the season. The base of the flowering axis does not become swollen
or form a new tuber, but a bud on the old parent tuber, in the axil of the lowest
sheathing leaf of the flowering shoot, provides for the continuity of the plant.
There is probably just a sufficient food reserve left in the old and somewhat
shrivelled parent tuber to enable the bud to commence its growth at the
beginning of the next season.
The next stage in development might well be a transition from the perennial
structure just described to the annual type of corm which occurs in several
genera in the Iridoideae, such as Hexaglottis, Moraea, etc., through the storage of
food reserves in an internode of the axillary bud and the complete shrinking
and almost entire disappearance of the parent corm or tuber.
4. The single internode corms of Hexaglottis, Homeria and Moraea
In the corms of the Iridoideae, as in the rhizomes and corms throughout the
family, the branching is sympodial and every year the terminal bud gives rise
to an aerial shoot while an axillary bud provides for the continuity of the plant.
The axillary buds of the members of the Iridoideae which have been examined
are morphologically not quite the same as those of the majority of the Ixioideae,
nor do they develop in the same way. With only one or two exceptions the type
of corm which occurs in the Iridoideae is associated with a dorsiventral leaf,
whereas in the Ixioideae the leaves are the isobilateral equitant type.
In most of the genera in the Ixioideae the food reserve is accumulated in two
or three or more abbreviated internodes at the base of the aerial axis and the
axillary bud which gives rise to the next season's shoot only develops the
following season, but in genera in the Iridoideae such as Moraea, Homeria, Hexa-
glottis and one or two others, the whole of the aerial axis dies away right down
to the base at the end of each flowering season and the food reserve is stored in
an internode of the axillary bud which develops at the same time as the aerial
axis, though the terminal growing point of the bud remains dormant until the
following growing-season. The growing point of the bud is often protected by
two or three very small scale leaves in which strong fibrous strands are usually
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 75
developed at an early stage, and the whole bud is generally enclosed within
a single persistent fibrous scale leaf which arises immediately below the swollen
internode. Between this leaf and the aerial axis there are usually a pair of
strongly keeled united prophylls which have no fibres and do not envelop the
bud except perhaps before it starts to swell. The whole structure, i.e. the base
of the aerial axis and the new axillary corm, is enclosed within another fibrous
scale leaf, the subtending leaf of the axillary bud, and outside this is a thin,
membranous sheath which extends up for a short distance and encircles the base
of the aerial axis. In the Ixioideae, as in the Iridoideae, the first leaf at the base of
the corm is nearly always a membranous sheath, either pale golden-brown or
colourless. The fibrous scale leaves outside this belong to the previous season's
corm.
Some of the corms of the Iridoideae are illustrated on pi. XIX. Fig. i is a
slightly enlarged longitudinal section of the old and new corms of Gynandriris
setifolia (L. f.) Foster, with the old outer corm tunics removed. At this stage,
about two months before flowering, the old corm is rather shrivelled and reduced
to half its original size as much of its food reserve has been used up. In this, as
in all these single internode corms, there is little if any trace of a stele. The
terminal bud of the old corm has developed into a shoot with a terminal aerial
axis (i) and the terminal bud of the next season's aerial shoot (viii) is already
formed at the apex of the swollen internode of the axillary corm-bud (ii) . As
this new corm continues to swell the base of the aerial stem is forced outwards
and becomes curved. At the base of the shoot are several fine adventitious roots
and above these the first two leaves of the shoot. The outer one (hi) is sterile.
It is very thin and colourless and soon becomes membranous. The inner one
(iv), which subtends the axillary corm-bud, is already firm and its fibrous
strands are well developed. The first leaves of the axillary bud are the united
prophylls (vi) which are not shown in this figure but can be seen in a transverse
section of the corm in fig. i a. Above the prophylls, shown in both figures, is the
first scale leaf of the axillary shoot (vii), which envelops the new corm and is
sterile.
The first leaf borne at the base of the aerial axis above the corm-bud is a short,
thin, membranous sheath which is not shown in either of the figures as it was
not observed in this early stage, though it was present in all the plants examined
about two months later. The leaf (v) shown in figs, i and la near the base of
the aerial axis is the first green leaf of the shoot and is of the dorsiventral type.
The internode below this leaf lengthens as the flowering axis develops and in
mature plants the node of the first green leaf is at or near the surface of the
ground.
Fig. 2 is a corm of Hexaglottis flexuosa with the outer scale leaf removed. At
the time when this was examined the plant was flowering and at this stage all
that remained of the previous season's corm was a very small, flat disc at the
base. The membranous remains of the prophylls can be seen between the aerial
axis and the new corm. These were removed and are shown in fig. 2a. Fig. 2b
Ann. S. Afr. Mus. Vol. XL.
Plate XIX
i, Gynandriris setifolia (L. f. ) Foster, longitudinal section of old and new corms, X 2; la, transverse
section of new corm, X2; i, aerial axis; ii, new corm; iii, membranous sheath; iv, lignified
scale leaf; v, base of first green leaf ; vi, prophylls; vii, first scale leaf of new corm; viii, growing
point of new corm. 2, Hexaglottis Jlexuosa (L. f.) Sweet, corm with outer tunics removed, xf;
i, prophylls; 2a, prophylls, x i£; 2b, 1. s. of corm; 2c, part of scale leaf, X f . 3, Moraea edulis
(L. f.) Ker, part of old scale leaf, x i-|-. 4, M. bituminosa (L. f.) Ker, 1. s. of corm, X i|; i, aerial
axis; ii, corm partly adnate to axis; iii, lignified base of first leaf ; 4a, base of leaf with cormlet
attached, x i£; 46, same, x f ; i, brown, viscid lining; 4c, part of old outer tunic with cormlet
attached. 5, Lapeyrousia corymbosa (L.) Ker, corm, xf ; 5a, flat base of corm, x f . 6-7, Romulea
bulbocodioides (De la R.) Baker. 6, corm, X i£; 6a, same, xf; 6b, corm with outer tunics
removed, xi{; i, lignified base of first green leaf; 6c, same corm with base of leaf removed,
xf; i, bud of next season's shoot. 7, new corm starting to develop, X i£; i, base of leaf;
ja, 1. s. of same corms, X 2\\ i, membranous sheath; ii, lignified scale leaf.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 77
is a longitudinal section of the corm and in this it can be seen that the base of
the aerial axis has been pushed outwards by the new corm so that, although it
is terminal and the corm axillary, the positions appear to be reversed. The first
leaf shown on the aerial axis is the membranous sheath which was not shown on
the younger axis of Gynandriris setifolia.
The only difference between the corm of Hexaglottis flexuosa and that of
Gynandriris setifolia is in the character of the fibrous tunics. In the latter they
are pale in colour and more or less of the 'herring-bone' type which is found in
several species of Moraea (see fig. 3, a portion of a scale leaf of M. edulis). In
Hexaglottis the fibres are much coarser and almost black. A portion of a corm-
tunic of H. flexuosa is shown in fig. 2c.
The corm of Moraea bituminosa differs in two or three respects from those of
Gynandriris and Hexaglottis and most other species of Moraea, although funda-
mentally it is the same type of corm. One difference is that all the scale leaves
in this species are not sterile but each subtends an axillary cormlet borne on a
short pedicel. A longitudinal section of a corm is shown in fig. 4 (pi. XIX), with
the old outer corm tunics removed. In M. bituminosa the leaf which appears to
subtend the new corm-bud is not a short scale leaf, as in Gynandriris and Hexa-
glottis, but though the base is fibrous and resembles a scale leaf, it is the first
long, green, dorsi ventral leaf of the plant. When removed from the axis it was
found that it does not subtend the new corm, but at its base is attached to the
pedicel of a small axillary cormlet. Two views of this are shown in figs. \a and
4#. (The closely allied species, M. uiscaria, has several of these cormlets with
more or less united pedicels.) One of these axillary cormlets still attached to
one of the old outer corm tunics is shown in fig. \c.
The axillary bud which has developed into the new corm appears to be
without a subtending leaf, but within the base of the dorsiventral leaf below is
a brown, viscid lining and, towards the top of the wide basal portion which
enclosed the corm, there are distinct traces of an inner skin. This is shown in
figs. 4# and aJj. The explanation for this somewhat anomalous structure appears
to be as follows : When the apical bud starts to develop at the commencement
of the growing-period cormlets borne on short pedicels arise in the axils of the
short scale leaves which have protected the apical bud during the dormant
period. The first leaf which develops at the base of the axis also subtends a
cormlet borne on a short pedicel which is partly adnate to the base of the leaf.
These cormlets might or might not develop and give rise to new plants at a later
date. From the number of old cormlets found among the old scale leaves which
envelop the corm it seems that many of them do not develop.
The bud which forms the new corm and which will give rise to the following
season's aerial shoot has probably arisen in the axil of one of a pair of united
prophylls, such as were found in the corms of Hexaglottis and Gynandriris, though
not in Moraea bituminosa. It is possible that such prophylls were present originally
but that they became completely detached as the corm started to swell and
disintegrated into the brown viscid substance lining the base of the leaf below.
78 ANNALS OF THE SOUTH AFRICAN MUSEUM
On the aerial axis of M. bituminosa there are nearly always two branchlets at
each node (occasionally three), apparently arising in the axil of a single sheathing
leaf, though when this is removed it can be seen that the second branchlet (and
the third when present) is subtended by one of a pair of brown membranous
prophylls.
Although the first cormlets formed are on short stalks or pedicels, the new
corm itself is sessile and, as can be seen in fig. 4, it is partly adnate to the base of
the aerial axis, yet another example of the various fusions and adnations which
occur in the family. The new corm has no sterile enveloping scale leaf at the
base, as in Gynandriris and Hexaglotiis; in other words it is the first internode of
the axillary bud which forms the corm, the only leaves which it bears being the
very small scale leaves at the apex which protect the apical growing point. The
three cormlets seen at the base in fig. 4 probably arose in the axils of the small
apical scale leaves of the previous season's corm.
5. The flat- based corms of Lapeyrousia, Romulea and Hesperantha
In the majority of genera in the Iridaceae the corm is more or less globose
(sometimes depressed-globose), but in several species of Hesperantha and Romulea,
one or two species of Geissorrhiza, and in all the species of Lapeyrousia in the
subgenera Sophronia and Ovieda, they have a very distinct and characteristic
shape which Dinter, in describing the corm of Lapeyrousia Vaupeliana, called
'broadly bell-shaped'. The scale leaves have a flat, circular base and articulated
to it is a broadly conical upper part.
A corm of Lapeyrousia corymbosa is shown in fig. 5 on pi. XIX, and its flat, solid,
fibrous base in fig. $a. The circular bases of the old scale leaves become detached
and eventually disappear with the remains of the old corms, while the upper
parts, of which the fibres are usually solid and regularly notched at the base and
coarsely and closely reticulate above, are persistent and remain as an outer
covering on the old corm. A succession of the upper portions of several old
scale leaves can be seen on the corm in fig. 5.
In Lapeyrousia the corms themselves, in the species of the subgenera Sophronia
and Ovieda which have been examined, are of the same type as those of Gynan-
driris and other members of the Iridoideae, that is, they are formed from a swollen
internode of an axillary bud which develops simultaneously with the aerial axis.
They differ only in shape and in having a small, well-defined stele. In these two
subgenera the node of the first green leaf, which is the isobilateral and not the
dorsiventral type, is on the aerial axis at or near to the surface of the ground, as
in Gynandriris, Hexaglottis and most species of Moraea and Homeria.
In the third subgenus of Lapeyrousia (Anomatheca) the corm is of the type most
common in the Ixioideae, with coarse or fine reticulate, fibrous tunics and no
distinct flat disc at the base, though in one or two specimens examined the corm
is slightly flat at the base. There are two or three basal leaves which arise from
nodes on the corm itself, their lignified bases forming some of the fibrous tunics
which envelop the corm, and the new growth commences the following season
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 79
from a bud in the axil of the upper of these leaves. This subgenus has diverged
so far from the other two subgenera that it should perhaps be treated as a
separate genus. Its affinity is with Freesia rather than with the other two sub-
genera of Lapeyrousia.
In Romulea triflora (Burm.) N. E. Brown and two or three other species the
corm is similar in shape to that of Lapeyrousia corymbosa but the scale leaves are
different, being solid and smooth, and the corm itself is of the Ixioideae type.
The first green leaf of the plant arises from a node on the corm which is the
swollen base of the aerial axis, and its axillary bud does not develop at all until
the following season. In several species of Romulea the corm is slightly or distinctly
obliquely flattened at the base. There is a small or sometimes fairly large, flat,
and somewhat rounded projecting portion on one side of the outer scale leaf,
and the upper part of the solid concentric corm tunic is very distinctly articu-
lated to the flat basal part in this region. The corm of R. bulbocodioides, which
is of this type, is shown in figs. 6 and 6a on pi. XIX. Fig. 6b is the same corm
with the outer tunics removed, showing the fibrous base of the first green leaf,
and fig. 6c is the same but with the base of the leaf removed to show the base
of the aerial deciduous part of the axis and the bud of the next season's
shoot.
At the commencement of the next growing period roots arise from the lateral
slightly bulging portion of the corm on one side of the axis, as can be seen in
figs. 7 and 7<2 (pi. XIX). Fig. 7 shows the old corm with the new corm starting
to develop. The old outer tunics have been removed and the small fibrous scale
leaf above the new corm is the lignified base of the first green leaf of the old
corm, removed from the lower corm and carried up by the new corm as it
starts to develop. Fig. ja is a longitudinal section, slightly enlarged, of the two
corms. The first leaf which envelops the new corm is a membranous, colourless
sheath. It has been mentioned before that the outer sheath of the corm seems
almost invariably to be membranous.
In many of the species of Lapeyrousia and Hesperantha which have been examined
the roots arise only on one side of the axis, growing out between the flat base of
the corm tunic and the region where the upper part is articulated to it (see
L. corymbosa, fig. 5) . The one-sided arrangement of the roots in these species
seems to indicate that these corms have originated fairly recently from an
underground axis which grew in a horizontal direction, and they probably
represent another case of a transition from a rhizome to a corm. As has been
mentioned earlier, there is a very close affinity between Schizostylis and Hesperan-
tha, the only difference between these two genera being that the former has a
rhizome and the latter a corm. Schizostylis probably represents the final stages
of a rhizome, in which the aerial portions of the plant are no longer perennial
but annual, and the species of Hesperantha with a flat-based corm represent the
first stage in a transition from this type of rhizome to a corm. The next stage
seems to be the disappearance of the flat base and a transition to the more or
less globose type of corm which is typical of the Ixioideae, in which the lignified
80 ANNALS OF THE SOUTH AFRICAN MUSEUM
bases of the green leaves arising from the corm form some of the protective
fibrous tunics of the corm.
In Romulea bulbucodioides the base of the first green leaf, which arises from the
corm, is lignified and similar to the upper portion of the outer scale leaf of the
corm. The green upper part of this leaf dies away with all the aerial parts of
the plant at the end of the season, but the lignified basal portion is persistent on
the corm throughout the dormant period, and finally becomes detached from
the corm by a clean-cut absciss layer at the base when the new bud starts to
develop (see figs. 6b, 6^ and 7 on pi. XIX).
In many species of Romulea, such as R. rosea and its numerous allies, the corm
is almost globose, with solid concentric tunics, and only a slight trace remains
of the fiat basal disc which is so conspicuous in R. bulbocodioides. In these species
there are two or three basal leaves arising from the corm and the aerial portion
of the axis is extremely abbreviated, usually only a few millimetres long and not
reaching to the surface of the ground. The same reduction of the basal disc has
taken place in some species of Hesperantha and the closely allied genus Geissorrhiza.
In describing the corms of the latter Foster stated that they are more or less
globose, or ovoid, or even conic in shape, always with a flattened base although
in some this is barely perceptible.
The flat basal part of the corm tunics in the corms described above might be
interpreted as the leaf base, the upper part which is articulated to it being the
lignified base of a rudimentary petiole. A reduction of the leaf base takes place
until it almost, or finally completely, disappears, and the lignified base of the
petiole is sessile on the axis. An analogy might' be found in the leaf bases of the
three shrubby genera which have been discussed in Part I of the paper. In
Klattia partita there is a conspicuous leaf base which projects above the node and
is persistent on the axis after the leaf has fallen (see pi. II, fig. 5) ; in Witsenia
mama the leaf base is reduced, especially in the dorsal region (see pi. Ill, fig. 3) ;
finally in Nivenia Stokoei (and other species of Nivenia) only a slight trace of a
leaf base remains on the axis after the leaf has fallen. (Fig. 7, pi. II, does not
show this as the leaves were cut offjust above the base to show the buds.)
In several species in the subgenus Ovieda of Lapeyrousia the flat-based corm is
associated with a more primitive type of inflorescence (see Part II of this paper),
and there is other evidence to suggest that the flat-based type of corm found in
the four genera mentioned above is a fairly primitive one. There are distinct
indications in some of the species in these genera that it develops into the globose
type of corm which is typical of the great majority of genera and species in the
Ixioideae. In many genera in the Ixioideae, however, the corm has probably
evolved along different lines, through an earlier transition from the rhizome
with evergreen leaves, to a perennial corm with evergreen leaves such as those
of Pillansia and Dierama, which have already been discussed, and from this to
the typical annual type of corm which is formed during one season, then passes
through a resting-period and dies away at the beginning of the next season as
its food reserves are used up.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 01
Lapeyrousia, Hesperantha and Geissorrhiza have been placed in the Ixioideae (or
Ixieae) in the various systems of classification, but Romulea, on account of its
abbreviated aerial axis and different type of inflorescence, was placed in a
different subfamily or tribe by the various botanists who have worked on the
family (i.e. in Sisyrinchieae by Bentham and Hooker and others ; in Crocoideae by
Pax and others). When describing the leaves of some species of Romulea Arber
suggested that the affinity was with some genera in the Ixioideae, such as
Hesperantha and Geissorrhiza, and in the structure of the corm as well as the leaves,
also in many aspects of the inflorescence and flower, there is evidence of a
distinct affinity with some of the genera in the Ixioideae. The leaf itself is of the
phyllode type which, according to Arber, is elaborated by secondary differentia-
tion from the fundamental flat equitant type.
6. The spinous roots of Moraea ramosissima
The corms of Moraea ramosissima (L. f.) Druce (M. ramosa Ker) appear to
differ so markedly from the typical Moraea corms already described that they
deserve some mention here. The peculiar spinous roots borne on the subterra-
nean axis of this plant have been described by Scott and were also discussed by
Arber (the observations of the latter based on herbarium material), but as more
material was at hand for examination than was available to either of them, one
or two features were observed which were not recorded by Scott or Arber.
This species, the largest and most ramified in the genus, is geophytic and
grows in fairly moist, often semi-shaded places ; it is sometimes abundant after
fires in really damp situations, such as stream-sides or slightly marshy ground.
The underground stem of one of the youngest plants examined is illustrated in
fig. i on pi. XX. This probably represents the first season's growth but it is
not known whether this plant originated from a seed or cormlet, though it was
most probably from the latter. A puzzling aspect about it is the length of the
vertical underground axis. There are no contractile roots to account for it,
and in older plants the axis is considerably shorter and the dense mass of cormlets
and roots is situated just below the surface of the ground, as can be seen in
figs, i a and lb.
The subterranean axis of the young plant consists of a series of nodes and fairly
long internodes, with a cluster of axillary cormlets at each of the nodes, though
only the uppermost, which is at the surface of the ground, is in the axil of a
subtending leaf. At the base of the first two nodes are ordinary adventitious
roots but at the third node there are a few normal adventitious roots and
between them and the cormlets three spinous roots have started to develop,
growing in an upwards direction. At the fourth node there are no ordinary-
roots but a single spinous root has started to develop.
Fig. i a shows the lower part of the axis of a slightly older plant which did not
produce any flowers and is probably the second season's growth. At the base
are several ordinary adventitious roots, but near the middle of the dense mass
of cormlets is a distinct whorl of spinous roots, the majority of them growing
Ann. S. Afr. Mus. Vol. XL.
Plate XX
1-3, Moraea ramosissima (L. f.) Druce. 1, underground stem with cormlets, first season, xf;
ia, same, second season, xf; lb, longitudinal section of same, third season (flowering), xf;
i, spinous roots; ii, normal adventitious roots. 2, longitudinal section of large cormlet, X i£;
i, membranous sheath; ii, lignified scale leaf; hi, inner lignified layer partly adnate to scale
leaf. 3, 3a and 3^, remains of membranous sheaths with spinous roots, x f . 4-5, M. plwnaria
(Thunb.) Ker. 4, Corm, xf; 4a, part of inner fibrous layer of scale leaf, X i£; 4b, cormlets,
5, part of inner and outer fibrous layers of scale leaf,
5a, 1. s. of cormlet, X 2. 6, Chas-
X I
manthe Jioribunda (Salisb.) N. E. Br., corm with leaves removed, from above, X f ; 6a, 1. s. of same
corm, xf; i, scars of leaves; ii, bud.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 83
upwards though one of them is growing downwards. Near the top of the mass
of cormlets is one which is considerably larger than the rest and it is almost
certainly the corm which will give rise to the next season's shoot. At the two
nodes above are leaves with their sheathing bases ruptured by the pressure of the
numerous axillary cormlets.
A longitudinal section of the base of a mature flowering plant is shown in
fig. ib. The first green leaf, which is the dorsiventral type, arises from the axis
just below the ground and above it is a succession of leaves, each subtending a
short axillary branchlet bearing numerous cormlets. The branchlets arising
from the abbreviated portion of the axis below the ground do not appear to have
subtending leaves but in the position they should occupy are the stout
spinous roots. The majority of these roots are at first negatively geotropic but
when they have reached a certain level, just below the surface of the ground,
they curve over and become positively geotropic.
There are two cormlets which are considerably larger than all the others and
probably one or both of these will give rise to a new flowering shoot the following
season. Fig. 2 is a longitudinal section of one of them. The outer sheath which
envelops the cormlet is thin, membranous and colourless. These thin outer
sheaths have either no visible vascular strands or else very fine strands,
apparently without any sclerenchyma, and soon break up and disappear. The
second scale leaf of the cormlet is firm and lignified but this also breaks up when
the corm begins to develop and either disappears or in some cases parts of it
remain and can be seen mixed up with the spinous roots.
The spinous roots were described by Scott, who stated that the only difference
of importance between them and the normal adventitious roots is that the more
internal vessels of the spinous roots are smaller. Arber suggested that the
association of one or more of the groups of cormlets with the basket-work of
interwoven spinous roots was probably fortuitous but it is doubtful whether this
view is correct. In all the specimens examined these spinous roots arise
immediately below the clusters of cormlets, and the position occupied by the
spinous roots and the apparent absence of subtending leaves to the underground
branchlets seems to have some significance. It is possible that the subterranean
clusters of cormlets originally arose in the axils of thin scale leaves, but that the
soft tissues between the vascular strands became ruptured and disappeared at
an early stage, although the vascular strands derived from the axis cylinder
remained and continued to grow. In one or two of the specimens examined the
membranous remains of leaves or sheaths were found which seem to lend some
support to this view. These are shown in figs. 3, 3a and 36. In all of them the
spinous roots appeared to be growing between two membranous skins. As has
been mentioned above, the lower green aerial leaves of the younger plants
become ruptured at the base as the cormlets develop. These can be seen in
figs. 1 and \a.
The fact that the spinous roots, as they start to develop, are almost always
negatively geotropic seems to lend further support to this view. As they no
84 ANNALS OF THE SOUTH AFRICAN MUSEUM
longer function as leaf veins it is possible that they have undergone various
modifications, both internally and externally, and the upper extremities of these
roots when they develop are usually positively geotropic and appear to function
as normal adventitious roots, which they frequently resemble in appearance
(see fig. 3) . The lower parts nearer to the axis have developed sharp spines and
the function of this portion is probably to form a protective basket-work
surrounding the cormlets, just as the lignified strands of the outer scale leaves
which persist after the softer tissues have died away, do in the case of the other
species of Moraea. Unfortunately an attempt made to cultivate some of the
cormlets for observation was not successful, so that up to the present it has not
been possible to observe any of the very early stages in the development of the
shoots.
There is a great deal of variation in the size of the plants in M. ramosissima,
as well as in the size of the basal clusters of cormlets and development of the
spinous roots. In many of the herbarium specimens examined the spinous roots
are fairly short and all are negatively geotropic. The numerous small cormlets
and larger cormlets or corms in M. ramosissima are all of the single internode type
which is found in other species of Moraea, Hexaglottis, etc.
7. The possible ligular origin of the inner part of the scale leaf of Moraea plumaria
The scale leaves of Moraea plumaria (Thunb.) Ker are unusual in the genus,
in fact in the family, and seem to lend some support to the hypothesis put forward
in the first part of this paper that in the Iridaceae there is an axillary stipular
outgrowth from the leaf base to which the base of the leaf is almost invariably
more or less adnate. In Moraea plumaria there appear to be two entirely different
types of scale leaf on the same corm, the outer a basket-like arrangement of a
few coarse reticulate fibrous strands (see pi. XX, fig. 4) and inside this a second
very different type with wider and more solid fibrous strands, which is shown in
fig. 4<7. In the older corms, between the inner layer and the corm itself, there
are sometimes clusters of axillary cormlets borne on short stalks which usually
force their way between the two layers, as can be seen in fig. 4. Some of these
cormlets, slightly enlarged, are shown in fig. 46.
At first the inner and outer fibrous layers are connected by a layer of slightly
fleshy tissue but these thin-walled cells soon break up and completely disappear,
leaving the inner fibrous layer attached to the outer only at the base, and
appearing like an appendage of the outer. This is illustrated in fig. 5, a longi-
tudinal section of a young corm with the corm itself removed so that only the
two fibrous layers are shown. A section of one of the cormlets is shown in fig. 5a.
In this last figure it can be seen that the axis extends above the first apparently
double scale leaf and there is an internode between this and the corm which is
enclosed in a brown membranous sheath. That the lower leaf is a single scale
leaf there can be no doubt, and the inner of the two fibrous layers might be
interpreted as an 'axillary' stipule or ligule which, in this species, is lignified.
The section of the young corm of M. ramosissima (pi. XX, fig. 2) shows a
Ann. S. Afr. Mus. Vol. XL.
Plate XXI
86 ANNALS OF THE SOUTH AFRICAN MUSEUM
somewhat similar structure, but in this case the inner layer is completely united
with the outer layer of the scale leaf for two-thirds of its length and is only free
in the upper third.
8. Relationships of the tribes considered mainly in connection with the underground organs
The various types of underground stems found in the South African Iridaceae
which have been mentioned in the preceding pages are arranged in tabular
form on pi. XXI, with the genera possessing these different types of organ
placed in the columns below. In drawing up this table one other factor has been
taken into consideration, that is, whether the inflorescence is spicate or not.
The great importance attached to the type of inflorescence in this family has
been discussed in Part II of this paper. It is necessary to consider the inflorescence
in conjunction with the subterranean parts of the stem as the changes and
reductions which have taken place in the aerial parts of the plant, which finally
led to the development of the spicate inflorescence in the Ixioideae, are associated
with changes in the underground parts of the stem, although this portion of the
plant has probably been the last to be affected by these changes.
Just as a few intermediate or transition stages exist which provide some clues
to the lines of development in the aerial parts of the plant, so also are a few still
found in the subterranean organs. In the genera Pillansia and Lapeyrousia the
intermediate conditions observed in the inflorescences are correlated with what
are considered to be transition stages in the subterranean axis. All the evidence
provided by the aerial and subterranean parts of the plants indicates that the
most primitive members of this family were perennial much-branched evergreen
herbs, or possibly small shrubs, with erect aerial stems arising from swollen basal
stems which developed in a horizontal direction on or just below the surface of
the ground.
In the diagram (pi. XXI) the types of underground stems have been divided
into eight groups, four of them having aerial leaves which are persistent for more
than one season, and four in which the aerial leaves are deciduous. Of these
eight groups, four are considered to represent distinct transition stages (see
columns 3, 4, 6 and 7). The rhizome with evergreen leaves is regarded as being
the most primitive type of subterranean stem and the others are believed to have
evolved from this.
Aristea is considered to be the most primitive genus found in South Africa.
The next is Bobartia, especially the two branched species, B. lilacina Lewis and
B. paniculata Lewis, which are mentioned in Part II. In one or two of the species
of Bobartia with a very much reduced inflorescence there is distinct evidence of
the rhizome undergoing a transition to a corm. This genus has therefore been
placed in two of the columns.
Dietes, although it still has a rhizome and evergreen leaves, and an inflores-
cence similar in some respects to that of Bobartia lilacina, shows a considerable
advance in the structure of the flower, which is like that of Moraea. There is
undoubtedly a close affinity between Dietes and Moraea and in the diagram this
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 87
is indicated by a dotted arrow. The type of corm found in Moraea and its allied
genera in column 5 has been described. It is the same in all the genera in this
column and in all of them it is associated with a dorsiventral leaf. No actual
intermediate conditions between a rhizome and a corm have been observed in
the large genus Moraea but it has been suggested that the type of development
in the 'tuberous corm' oiFerraria might indicate the lines along which the corm
of Moraea, etc., has evolved.
All these genera, i.e. Aristea, Bobartia and Dietes in column 1, Bobartia and
Ferraria in column 3, and Hexaglottis, Homeria, Moraea, Gynandriris and Galaxia
in column 5, belong to the Iridoideae. (According to Pax, Galaxia belongs to the
Crocoideae; the reasons for including it with the Iridoideae will be given in the
final part of this paper.) The limits of this tribe, number 1, are shown by the
double lines in the diagram. This is regarded as being the most primitive tribe
of the Iridaceae. In this tribe, though some of the genera have a rhizome and
others a corm, there is less evidence of the transition from the former to the
latter than there is in the more advanced Ixioideae.
The origin of the three shrubby genera in the second column has been the
subject of a good deal of discussion, Marloth and a few other botanists regarding
them as being of exceedingly ancient origin and possibly representing relics of
an older flora. They exhibit certain features, however, which are more
advanced than those of Aristea, as in the inflorescences which in the shrubby
genera give some evidence of having undergone further reductions than those
of some of the more primitive species of Aristea, as well as in the sessile flowers,
all with a perianth tube (in Klattia very short). They form a small but very
distinct group which Weimarck placed in a separate tribe, Nivenieae (together
with three genera found in South America and one in Australia). I agree with
Weimarck in regarding these shrubby genera as constituting a distinct tribe and
consider that this tribe, which is confined to the Southern hemisphere, is inter-
mediate between the Iridoideae and Ixioideae, though nearer to the former.
It is very possible that at one time this tribe was considerably larger than it is
to-day. The three South African genera, although they have very similar
subterranean stems and leaves, are quite distinct from each other, with very
different flowers which are arranged in various types of reduced complex
inflorescences. Their development has probably been greatly influenced by
their habitat; the majority of the comparatively few species which occur in
South Africa (one in Witsenia, two in Klattia and eight in JVivenia) grow in moist
situations, mostly along stream-sides or in marshy places, so that on the whole
they have not been as greatly affected by seasonal changes in moisture and
temperature as nearly all the other genera in the Iridoideae and Ixioideae. It is
possible that some of the more primitive features have survived in the vegetative
parts of these plants and their characteristic swollen, more or less subterranean
stems might even represent a more primitive type of subterranean axis than the
rhizome. Adamson, who studied the stems of the shrubby genera, noted that
although secondary thickening takes place in the swollen basal stems of these
88 ANNALS OF THE SOUTH AFRICAN MUSEUM
plants, as in their aerial stems, there is no sign of it in the rhizomes of some of
the species of Aristea which he examined.
The Ixioideae (No. Ill in the diagram) is the most advanced tribe and is by
far the largest. As will be seen in the diagram, the great majority of the genera
fall into the last column, in which the more or less globose corm is formed
through the accumulation of food reserves in the abbreviated subterranean
portion of the aerial axis. In these genera in column 8 the inflorescence is
spicate (except in Romulea, Syringodea and Crocus — the last not a South African
genus) , and in all of them the aerial part of the axis as well as the aerial parts
of the basal leaves arising from the corm disappear at the end of the flowering-
season.
There are a few exceptions in the Ixioideae which provide some distinct clues
as to the lines of development in this large group. One of the most important is
the monotypic genus Pillansia. The inflorescence of this genus was discussed in
Part II. Although the inflorescence is not spicate, but more or less intermediate
between the type of inflorescence found in one or two species of Nivenia and a
spicate inflorescence, and the subterranean stem is intermediate between a
rhizome and corm, Pillansia cannot rightly be included with the Iridoideae, nor
with the Nivenieae, and is best placed in the Ixioideae. It represents the most
primitive genus in this tribe but is undoubtedly closely allied to Watsonia
(indicated by a dotted arrow) and almost certainly represents a survivor of the
ancestral type of that genus. There is some reason to believe that the corm of
Watsonia passed through an intermediate stage like that of Dierama, that is a
perennial corm with evergreen leaves.
The corm of Dierama has been discussed, and its transition to the annual type
found in D. trichorhizum. This genus has therefore been placed in two columns,
4 and 8, connected by an arrow, though all except one of the species belong in
column 4. The affinity between Dierama and Ixia (section Morphixid) has also
been mentioned; this is indicated by a dotted arrow.
The only genus in the Ixioideae with a rhizome is Schizostylis, though this
rhizome differs from those in the Iridoideae in having deciduous aerial leaves.
It has been suggested that the survival of this primitive type of subterranean
axis is probably due to the habitat, in wet situations, and that it represents an
intermediate condition, being the final stages of a direct transition from a
rhizome to a corm without passing through intermediate stages such as those in
Pillansia and Dierama. There is a very close affinity between Schizostylis and
Hesperantha (indicated by a dotted arrow), and the flat-based type of corm found
in Hesperantha, as well as the other genera in column 7, is also regarded as a
transition stage, which in turn passes into the more or less globose corm which
is typical of the great majority of the Ixioideae. As both types of corms are found
in Hesperantha this genus has been placed in columns 7 and 8, as are also
Geissorrhiza, Romulea and Lapeyrousia. Geissorrhiza is closely allied to Hesperantha,
and in this slightly more advanced genus there are only one or two species with
a distinct flat base, the majority of the species falling into the last column.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 89
The close affinity between Schizostylis and Hesperantha and the transition from
the rhizome of the former to the corm of the latter leads one to believe that the
flat-based corms oi Romulea and Lapeyrousia probably evolved along similar lines,
but that in these two genera the transition to a corm took place at a slightly
earlier stage, before the inflorescence had reached its final reduction to a spike,
whereas in Schizostylis the spicate inflorescence was evolved before the final
transition from a rhizome to a corm.
Pax's tribe Crocoideae is indicated by dotted lines, group l\a. This tribe,
according to Pax, consists of the genera Galaxia, Romulea, Syringodea and Crocus.
It is not proposed to discuss this group at present as it will be mentioned in more
detail in the next part of this paper; in my opinion it is an artificial one
consisting of one member of the Iridoideae {Galaxia) and three of the Ixioideae.
It will be noticed that Lapeyrousia has been placed in three positions. Only the
subgenus Anomatheca belongs in column 8, but, as pointed out before, in the
subgenus Ovieda there is more than one type of inflorescence, many of the species
having a simple or branched spicate inflorescence, while in others there is a
more primitive type, with intermediate stages between the two. All the genera
placed in column 7 should almost certainly be included with the Ixioideae. A
system of classification based on correlated morphological characters is suggested
in Part IV; for the present it is intended merely to point out that the type of
corm in Romulea and Syringodea is the same as that of a few genera in the Ixioideae,
and not like that of the Iridoideae.
Part IV. The Flowers and Systems of Classification
1. Introductory remarks
This final part is divided into two sections ; the first deals rather briefly with
the flowers and one or two controversial taxonomic groups and the second with
the phylogeny and classification. In the second section the correlated morpho-
logical characters of the vegetative and floral organs are considered in connection
with a proposal for an amended system of classification which is not entirely in
accordance with any of the previous systems.
In the introduction to this paper it was pointed out that a great deal of
confusion has resulted from relying too much on the morphology of the flowers
alone, without taking into consideration the morphological character of the
other parts of the plant. It is not intended to underestimate the value of the
floral characters for there is no doubt that they are of great importance to the
systematist, but in this family it is not always possible to differentiate the tribes
or genera merely by the shape or form of the flowers or their styles and stamens.
Where this was done in the past it resulted sometimes in an artificial grouping
of a number of species which actually belonged to two or three or more different
genera, as in the case of Antholyza, Acidanthera, Ixia and Tritonia.
It is felt that the floral characters of the Iridaceae are on the whole fairly
well known, since great importance has been attached to them in systematic
90 ANNALS OF THE SOUTH AFRICAN MUSEUM
work, and it is not proposed therefore to analyse in detail the morphological
features of the flowers, but only to discuss one or two of them, and to mention
a few of the anomalies which have been observed in the flowers of some of the
genera.
In the more primitive genera in the Iridoideae, though there is a great deal of
variation in the type of inflorescence in some of the genera, the flowers in the
various genera are as a rule fairly homogeneous, although variations do occur,
such as the occasional development of a perianth tube in a single species of a
genus which normally has completely free segments, also a tendency to a reduc-
tion of the inner whorl of the perianth segments in one or more species in one
or two of the genera. In the more advanced Ixioideae the inflorescence in almost
all of the genera is reduced to its simplest form, either a simple or branched
spike (except in Pillansia and some species of Lapeyrousia) , but a greater degree
of variation occurs in the flowers in some of the genera and it is this group, with
its intermediate forms and borderline species apparently connecting some of
the genera, which presents the most problems for the systematist.
In many of the larger genera there are almost invariably one or two species
in which the flowers do not conform with the description given of the genus,
although the vegetative characters are usually more or less typical. The flowers
of these genera sometimes cover a fairly wide range, varying from the simple,
more primitive forms to more highly evolved forms, and often provide important
phyletic indications of actual affinities as well as of evolutionary trends.
Section I. The Flowers
2. Symmetry
A character to which some importance has been attached in systematic work
is the symmetry of the flowers. The Iridoideae and Crocoideae present no problems
in this connection as the four genera placed in the Crocoideae by Pax all have
actinomorphic flowers and throughout the Iridoideae, with probably only one
exception, the flowers are also actinomorphic. The only genus which is known
to have zygomorphic flowers in this tribe is endemic in Australia (Diplarrhena) .
In the Ixioideae, however, both actinomorphic and medianly zygomorphic
flowers occur, and it has been found that in some of the genera previously
described as having either one or the other, both types occur though usually
one or the other predominates. For instance, in Lapeyrousia and Babiana both
types occur but in the majority of species in these genera the flowers are zygo-
morphic. The genus Babiana has been divided into two subgenera, the one with
subequal perianth lobes and more or less actinomorphic flowers, the other with
more or less unequal perianth lobes and zygomorphic flowers, but an examina-
tion of all the species in this genus has proved this arrangement too artificial to
be upheld as the first group merges into the second through intermediate stages.
In Hesperantha and Geissorrhiza nearly all the species have actinomorphic
flowers with equilateral stamens, but a few exceptions have been recorded in
Ann. S. Afr. Mus. Vol. XL.
Plate XXII
i, Watsonia marginata (L. f.) Ker, flower, front view, xf; la, perianth tube with upper half
opened to show base of filaments (i) and staminodes (ii), X i|; ib, part of perianth tube, side
view, xi|; i, base of filament; ii, staminode; iii, callus. 2, Gladiolus species (aff. G. trichonemi-
folius), flower laid open, xf; 2a, perianth tube with stamens and stigmas, xif; 2b, bract, xf;
2c, bracteoles, x f ; 2d, corm, x f. 3, Watsonia Pillansii L. Bolus, perianth tube opened to show
filaments and staminodes, X f ; 3«, part of perianth tube with base of filament (i) and staminode
(ii), Xi|. 4, Moraea Cooperi Baker, flower, xf; i, pedicel; ii, ovary; iii, perianth tube. 5,
Tritonia scillaris (L.) Bak., stamen, front view, X 3^; 5a, same, side view; i, opening. 6, Babiana
stricta (Ait.) Ker, stamen, front view, xi|; i, connective. 7, Exohebea lata (L. Bol.) Foster,
anther, front view, X 1 1; ya, same, back view. 8, E. parviflora (Jacq.) Foster, stamen, back view,
X i^; 8a, same, front view. 9, E. lata, style branches and stigmas, X i£; ga, same, from above.
10, Moraea plumaria (Thunb.) Ker, style branches and stigmas (i), x \\.
92 ANNALS OF THE SOUTH AFRICAN MUSEUM
which the stamens and style are more or less arched or all directed to one side,
as in Hesperantha grandiflora, Geissorrhiza rochensis and G. Lewisae. Although the
position of the style and stamens can be seen clearly in living flowers, it is often
obscured or lost in dried material, from which many of the original descriptions
were drawn.
The asymmetry of the stamens in flowers which are otherwise regular is
undoubtedly the first step towards the more highly evolved types of medianly
zygomorphic flowers in which the perianth tube is curved and the flowers tend
towards a horizontal position with the stamens and style arched under the
uppermost lobe which often becomes expanded and hooded, as in the great
majority of species of Gladiolus, or elongated, as in Chasmanthe, Anomalesia and
one or two other genera. In the more highly evolved flowers of these genera
the elongation of the uppermost perianth lobe is accomplished at the expense
of the other lobes which are nearly always very much reduced. This is apparent
in some species of Gladiolus, especially in the orchidiflorus group, as well as in
Antholyza, Anomalesia and Kentrosiphon. In Anomalesia the three lower lobes are
extremely reduced and remain curved inside the perianth tube (see pi. XXIII,
fig. 4) , while in Kentrosiphon a pouch has developed below the three lower lobes
and only a trace remains of the lowest lobe, the four lateral lobes being very
much reduced and only the conspicuous scarlet uppermost lobe well developed
(see pi. XXIII, fig. 5).
In most species of Watsonia, although the uppermost perianth lobe is usually
arched above the stamens, it is scarcely at all expanded or wider than the other
lobes. There are one or two species of Watsonia in which the flowers are very
nearly regular, as in W. marginata, where, except that the perianth tube is
slightly curved, the flowers are actinomorphic (see figs. 1 and \a on pi. XXII).
In Gladiolus, the genus with the greatest number of species and certainly the
greatest range of distribution (South and tropical Africa, Europe, and Asia),
there are in South Africa a few species in which the flowers have subequal
perianth lobes and are only very slightly zygomorphic, as well as one or two
in which they are definitely actinomorphic. Botanists have hesitated about
placing these anomalous species with actinomorphic flowers in the genus, yet
the corm, leaves and bracts, as well as the flattened alate seeds, are typical of
the genus, or rather of the particular group of species to which they are clearly
related within this large and heterogeneous genus.
One example which might be given is the plant known as Geissorrhiza
Patersoniae L. Bolus. Foster, in his revision of Geissorrhiza, followed L. Bolus in
treating this as a Geissorrhiza, but placed it in a separate subgenus. In my
opinion, however, although it obviously represents a link between Geissorrhiza
and Gladiolus, it should rather be placed in the latter genus near to G. edulis
and G. permeabilis, from which species it differs only in having actinomorphic
flowers. (Incidentally this species was first collected by Thunberg and named by
him Gladiolus elongatus. N. E. Brown upheld this species although he did not
associate it with Geissorrhiza Patersoniae.) Another borderline species which
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 93
seems to connect these two genera is one which has not so far been described.
The flowers are completely regular (see figs. 2 and 2a on pi. XXII), yet all its
other characters, such as the flat alate seeds, the bract, bracteoles and corm (see
figs. 2b, 2c and 2d) are typical of the group of Gladioli comprising the species
G. linearis, G. erectiflorus and G. trichonemifoliiis. If the flower alone were to be
used as a diagnostic characteristic this plant would be placed in the genus
Geissorrkiza, although the other characters do not conform with those of that
genus.
3. The perianth tube
Another floral character which should be used with discretion is the length
of the perianth tube. As was pointed out in a paper published by myself in 1941 ,
elongation of the perianth tube occurs in many genera, such as Lapeyrousia, Ixia,
Babiana, Gladiolus and even Romulea. A characteristic of the last-named genus
is the short, funnel-shaped perianth tube, yet two or three species with a long
perianth tube have been found in South Africa. Very often intermediate forms
connect the short and long-tubed forms so that this character alone is not a
satisfactory or sufficiently limiting factor to use in subdividing a tribe or genus.
Although the length of the perianth tube is of relative unimportance, the
presence or absence of a perianth tube is an important character. In the
Ixioideae the flowers are sessile and all have a perianth tube, even though in some
cases it is very short, as in Melasphaerula and a few species oiGeissorrhiza. In the
Crocoideae both Galaxia and Syringodea have very short pedicels and long perianth
tubes. In the South African members of the Iridoideae, excluding the shrubby
genera Klattia, Witsenia and Nivenia, the flowers are always more or less pedicel-
late and the perianth segments nearly always free or almost so. There are,
however, three anomalous species in this group in which the flowers, as well
as being distinctly pedicellate, have a short but quite distinct perianth tube.
These three species, belonging to three different genera in which the perianth
segments are described as free, have been the cause of some confusion owing to
the fact that, in the case of two of them at any rate, the original descriptions
were made from poorly dried herbarium material in which the flowers were
evidently not dissected, for no mention was made of the perianth tube. One of
these is Bobartia macrospatha Baker (syn. B. tubata Gillett), which has a pedicel
up to 5 mm. long and a perianth tube up to 1 cm. long. Gillett has pointed out
to me that the plant he described as B. tubata was later found to be the same as
Baker's B. macrospatha. He had dissected a flower of the type and found that it
had a perianth tube although Baker made no mention of it in the specific
description and the generic description in the Flora Capensis seemed to deny its
existence.
Another species, which necessitates some amendment to the generic descrip-
tion, is Hexaglottis virgata (Jacq.) Sweet. This has a pedicel about 5 mm. long
and also a perianth tube 5 mm. long, although in the generic description in the
Flora Capensis the perianth is described as 'cut down to the ovary'. In the Flora
94 ANNALS OF THE SOUTH AFRICAN MUSEUM
of the Cape Peninsula it is regretted that I also omitted to observe the presence of
a perianth tube in this species.
The third species is undoubtedly a Moraea although it appears to be quite
anomalous in that genus. At present it is known by three different names,
Moraea Cooperi Baker, Gynandriris stenocarpa (Schltr.) Foster {Moraea stenocarpa
Schltr.) and G. apetala (L. Bol.) Foster {Moraea apetala L. Bol.), but the three
type specimens have been examined and there is no doubt that they all represent
the same species; so that Baker's name, the first given to this species, must be
retained and the other two become synonyms. Baker's description is incorrect
and quite misleading. He described the flowers as 'lilac', whereas they are pale
yellow, though they usually turn mauvish when dried. He made no mention of
the perianth tube and described the inner perianth segments as 'oblanceolate',
but in this species there are no inner segments! Working with poorly dried
specimens it is very possible that he mistook one of the style crests for an inner
segment. He also described the spathes as two- to three-flowered, whereas they
are one-flowered, a character which is probably unique in the genus. In this
respect, as well as in having a perianth tube, Moraea Cooperi differs from all other
species in the genus.
The most complete description, from living specimens, was given by L. Bolus,
but after having examined a large number of living specimens it was found that
there is one point which requires clarification. In describing Moraea apetala in
1929 Dr. Bolus stated that the ovary is produced into a beak like a perianth tube.,
while in all the specimens examined by myself in 1950 it was found that there
was an articulation at the top of the ovary, below the short cylindrical perianth
tube which is continuous with the segments (see fig. 4 on pi. XXII). The genus
Gynandriris {Helixyra Salisb. ex N. E. Br.) is separated from Moraea on account
of its membranous spathes and the beak-like elongation above the ovary, with
the perianth segments articulated to its apex, so that if L. Bolus's description
were correct then this species should belong rather to the genus Gynandriris ; this
combination was actually made by Foster. In Moraea Cooperi the spathes are
not membranous, nor does the ovary extend up in a short, sterile projection, so
that this species cannot be included with Gynandriris but should remain in the
genus Moraea, where it should be placed in the subgenus Vieusseuxia which is
characterized by having the three inner perianth segments much smaller than
the outer, sometimes tricuspidate or, as in M. tripetala, extremely reduced and
filiform, sometimes barely perceptible. In M. Barnardii L. Bol. they are com-
pletely suppressed. M. Cooperi and M. Barnardii are the only two South African
species so far known which have only three perianth segments, with no trace
of the inner whorl. One of the characters distinguishing Moraea from Iris is the
presence of a perianth tube in Iris and the absence of one in Moraea, so that
M. Cooperi, with its short perianth tube, provides a link between these two
closely allied genera.
It was mentioned in Part II of this paper that the reduction from a compound
inflorescence to a simple spicate one appeared to be correlated with other
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 95
variations. One of these was that the reduction in the number of flowers
resulted in the production of a smaller number of flowers of very much firmer
texture, each individual having a flowering period of a few days instead of only
a few hours. Another variation to which reference was made was that the
reduction of the internodes of the branches and of the pedicels was sometimes
accompanied by an elongation of the ovary, as in the section Pseudaristea of
Aristea, or by the development of a perianth tube. In the three anomalous
members of the Iridoideae mentioned above considerable reductions have taken
place in each case. In Hexaglottis virgata the internodes of the rather numerous
lateral branches are completely or almost completely suppressed, so that the
flower-clusters or rhipidia (each two-flowered) are sessile on the axis and the
inflorescence has the appearance of a spike. In Moraea Cooperi and Bobartia
macrospatha the cymes are one-flowered, and in both, and Hexaglottis virgata,
the pedicel is considerably shorter than is usual in these genera, so that the
ovary, instead of being exserted, remains enclosed in the spathes or bracts.
In a discussion of the morphology of the genus Nivenia Weimarck made the
following statement : 'The tube of Nivenia flowers corresponds biologically in
some Aristea species to the pedicels (e.g. A. alata), in other species to the cylin-
drical or prismatic ovary (sect. Pseudaristea).' This interpretation of the perianth
tube is probably correct, and the slender, stalk-like sterile extension of the ovary
in Gynandriris, and observed also in one or two species of Ferraria, seems to
provide some evidence of a preliminary step towards the formation of the
perianth tube. It is very probable that this stage is followed by the transition of
the point of articulation from the top of the ; beak' to the top of the functioning
part of the ovary, so that the sterile upper part, with no articulation between it
and the segments, becomes a part of the perianth instead of an extension above
the ovary.
Part of the perianth tube in the Ixioideae, possibly the whole of the lower portion
of the tube up to the point where the stamens are inserted, was probably derived
in a similar manner from the pedicel, the upper part, which is present in many
of the genera, being formed by the fusion of the lower part of the perianth
segments. In some genera the stamens are inserted near the top of a slender
cylindrical tube (e.g. Exohebea, Thereianthus and several species of Ixia), in others
at or near the middle of the tube, as in Watsonia, Gladiolus, etc., but in either case
the portion of the tube below the stamens is slender and more or less erect. When
the tube is curved the bend is almost invariably at the junction of the upper and
lower parts, as in Watsonia and Gladiolus. In some cases the lower part passes
gradually into the upper part which is often funnel-shaped, as in some species
of Gladiolus ; in others the perianth tube is clearly differentiated into two distinct
portions, the upper either funnel-shaped, as in many species of Gladiolus, or
broadly cylindrical as in many species of Watsonia, also in Homoglossum,
Anapalina and Chasmanthe.
In two species of Chasmanthe, C. aethiopica and C. Peglerae, the differentiation
of the perianth tube into two distinct parts is further emphasized by the develop-
Ann. S. Afr. Mus. Vol. XL.
Plate XXIII
1-2, Chasmanthe aethiopica (L.) N. E. Br. i, spike, X f, arrows indicate direction of twist; \a, part
of perianth tube of bud, X 2 ; ib, part of perianth tube of open flower, with spiral twist in basal
portion, X2; ic, bract, xf; id, bracteoles, X f . 2, capsules, xf; ia, open capsule, xf;
lb, seed, X i-|; i, point of attachment; 2c, transverse section of seed, X i|; i, thick yellow testa.
3, Anapalina revolula (Burm.) N. E. Br., flower, first stage, x-f; 3a, same, second stage. 4,
Anomalesia Cunonia (L.) N. E. Br., flower, xf; i, three lower perianth lobes folded inside tube.
5, Kentrosiphon saccatus (Klatt) N. E. Br., flower, x f ; i, lowest perianth lobe.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 97
ment of three pouches at the base of the upper part. Nectar, secreted in the
ovary, passes up the short slender basal part of the perianth tube and collects
in these pouches. The stamens are inserted at the base of the upper part of the
perianth tube and the basal part becomes spirally twisted as the flowers open.
In this respect it behaves somewhat in the same manner as the pedicel and
ovary, or ovary alone, of some genera in the Orchidaceae. The secund position
of the fully open flowers in C. aethiopica is brought about by the twisting of the
basal part of the tube. The buds, as they first emerge from the axis, are more or
less distichously arranged but slightly twisted, one of the upper side lobes being
opposite the subtending bract. As the bud begins to open the basal part of the
tube begins to twist in towards the axis. This is shown in fig. i on pi. XXIII.
At one stage the half-open flowers face towards the axis and for a short time they
are inverted but the tube continues to twist and when the flowers are fully open
the perianth tube has more or less completed a spiral twist. The ovary and
lower half of the perianth of a bud, before the basal part of the tube has started
to twist, is shown in fig. \a, and the spiral twist in the tube of a mature flower
in fig. ib. In C.floribunda the flowers are more or less distichously arranged in
the spike and the perianth tube is less twisted than in C. aethiopica, and in this
species there is only one rather shallow pouch at the base of the broadly
cylindrical upper part of the tube.
4. The genus Chasmanthe
As there has been some difference of opinion about the status of Chasmanthe
and two or three of the other genera established by N. E. Brown twenty years
ago, it seems advisable to discuss them briefly here. Chasmanthe is one of the six
new genera described by Dr. Brown in 1932 when he divided up the species
which had previously been included in Antholyza. In a note on N. E. Brown's
subdivision of the genus Antholyza, published in 1941, E. P. Phillips commented
as follows:
'The writer considers that Brown has succeeded in clearing up much of the
prevailing confusion though he has gone too far in proposing so many genera.
In the writer's opinion the genera Petamenes Salisb., Kentrosiphon N. E. Br.,
Chasmanthe N. E. Br., and Anomalesia N. E. Br. should be grouped together under
the oldest name Petamenes Salisb. All these genera are characterized by the
elongated upper perianth-lobe which is differently shaped to the other five lobes
and are concave or hooded. This appears to be a very natural grouping, but
the subsidiary characters used by Brown to separate the genera, e.g. the saccate
perianth-tube of Kentrosiphon, the reflexed perianth-lobes of Anomalesia, and the
slight differences he gives between Petamenes and Chasmanthe do not warrant
generic status.'
The fallacy of relying entirely on floral characters for grouping species together
in this family, more especially in the Ixioideae, has already been pointed out in
this paper. In placing Chasmanthe with Petamenes, Kentrosiphon and Anomalesia,
Phillips has taken into consideration only the floral characters of these genera,
98 ANNALS OF THE SOUTH AFRICAN MUSEUM
which are somewhat similar. Hutchinson, in his Families of Flowering Plants,
published in 1 934, also grouped Chasmanthe with Petamenes though he maintained
both Anomalesia and Kentrosiphon as separate genera.
I cannot agree with Phillips or Hutchinson, and consider that Brown's genus
Chasmanthe, with the exception of three of the species which he placed in this
genus, is well defined and quite distinct from Petamenes. Two or three important
characters do not agree with the generic description of Petamenes given by-
Phillips. For instance, he described the corm as 'small, globose or sub-globose' ;
this applies to the corm of Petamenes but the corm of Chasmanthe is not small,
that of C. Jlorib unda being among the largest in the family, nor is it globose.
Two views of the somewhat flattened corm ofC.foribunda, with the corm tunics
removed, are illustrated on pi. XX, figs. 6 and 6a. The capsule Phillips described
as 'ellipsoid, shorter than the bracts', and the seeds as 'many, flat, broadly
winged', but again this applies to Petamenes only, for in Chasmanthe the capsule is
somewhat rounded and considerably longer than the rather short bract and
the few seeds produced are fairly large and oval, with a smooth, shining, thick
and brightly coloured testa. They are distributed by birds and not by wind,
as are the winged seeds of Petamenes. The capsule and seeds of C. aethiopica are
illustrated in figs. 2 and 2c on pi. XXIII. The testa in this species is bright
yellow or orange.
The three species which should be excluded from Chasmanthe are C. fucata
(Bak.) N. E. Br., C. cqffra (Bak.) N. E. Br. and C. intermedia (Bak.) N. E. Br.
Both Hutchinson and Phillips upheld Brown's genus Curtonus, but L. Bolus
considered that it should be combined with Chasmanthe, according to the follow-
ing note made by her in 1933 (S.A. Gard. and Country Life XXIII, p. 46):
'Another genus which rests upon too weak a basis to be stable is Curtonus,
containing only one species, C. paniculatus, and occurring in the Orange Free
State, Transvaal and Natal. It was placed by Baker next to what is now
Chasmanthe fucata, from the Khamiesberg (Namaqualand) , and there seems to be
no ground for upsetting this arrangement.' There is no doubt that Dr. Bolus
is correct in recognizing that these two species should not be separated, but in
my opinion they should be placed together in the genus Curtonus, which, although
closely allied to Chasmanthe, seems to be sufficiently distinct to merit generic
status.
The other two species, C. cqffra and C. intermedia, should be transferred to the
genus Anapalina. The corms, leaves, bracts, fruits and seeds of these two species
are all typical of the genus Anapalina and none of these organs are the same as
those of Chasmanthe. The corms are deep-seated, as in Anapalina, and enclosed in
wiry reddish-brown fibrous tunics which are not reticulate and extend up in a
long neck. These characters do not agree with those of Chasmanthe, though they
are typical of Anapalina and the closely allied genus Exohebea. The difference
between the leaves of Anapalina and Chasmanthe was pointed out in the first part
of this paper. The leaves of C. cqffra and C. intermedia are typical of the former
genus, with only a few prominent primary veins, instead of numerous veins and
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE 99
a pseudo-mid-rib as in Chasmanthe. The bracts and bracteoles of C. aethiopica
are shown in pi. XXIII, figs, ic and id; the bifid bracteoles are of the type most
commonly found in the Ixioideae. In C. cqffra and C. intermedia the bracteoles are
slightly longer than the bracts and they are of the type peculiar to Anapalina
and Exohebea, which were discussed in the first part of this paper. The fruits
and seeds also are quite unlike those of Chasmanthe, described above, and are
exactly like those of Anapalina. N. E. Brown's description of the seeds of Anapalina
was incorrect, as was pointed out by L. Bolus : 'Dr. Brown describes the seeds as
"flattened, winged all round" : those I have seen are four-sided, each face
being sculptured, or bearing processes, and being surrounded by a wing.'
It was only on account of the analogous flowers, with the uppermost lobe
much longer than the other five, that the two species just mentioned were
placed in Chasmanthe. In Anapalina the perianth lobes vary from subequal, as
in A. triticea, to distinctly unequal. In A. revoluta there is a clear indication that
the uppermost lobe tends to become elongated and in some of the forms of this
widespread and variable species it is considerably longer than the other lobes,
as can be seen in the flower illustrated in fig. 3 on pi. XXIII. In some specimens
collected in the Bredasdorp Division the uppermost lobe is even longer than in
the flower illustrated. The flowers are protandrous and a later stage with the
perianth lobes recurved is shown in fig. %a.
With regard to two of N. E. Brown's genera, Anomalesia and Kentrosiphon, I
am inclined to agree with Phillips in considering that Brown created too many
genera, and consider that Kentrosiphon and Anomalesia should be combined under
the latter name. The only difference between them is the presence of the pouch
or spur in Kentrosiphon and the reduction of the two upper lateral perianth lobes.
The corms, leaves, bracts, fruits and seeds of these two genera are the same.
On pi. XXIII, figs. 4 and 5, are shown a flower of Anomalesia cunonia and one of
Kentrosiphon saccatus. Whether Phillips is right in sinking both genera in Petamenes
I am not prepared to say until there has been an opportunity of examining
more material of Petamenes, but it is very likely that Anomalesia (including
Kentrosiphon) is sufficiently distinct to be maintained as a separate genus.
L. Bolus remarked of Petamenes abbreviatus, on which the genus was founded :
'This barely escapes being placed in Homoglossum and is one of the critical species
that are difficult to place : if it were not for the short side-sepals and oblique
tube-mouth it could be put into Homoglossum.' Homoglossum, in its turn, is
extremely closely allied to Gladiolus and in fact there are two or three records
of a natural hybrid between Homoglossum Priorii and Gladiolus maculatus.
The whole group of the most advanced members of the Iridaceae, which
Hutchinson placed together in the tribe Antholyzeae, presents more problems
than any other and requires further critical study. The tribe is actually rather
an artificial one based entirely on the fact that the flowers are, on the whole,
more zygomorphic than in the Gladioleae tribe, but it is very doubtful whether
there is sufficient justification for this distinction. This matter will be discussed
again in the final section.
100 ANNALS OF THE SOUTH AFRICAN MUSEUM
5. The stamens
Throughout the family the number of stamens is three, but occasionally in
abnormal flowers one or other of the stamens of the inner whorl is developed.
The genus Watsonia is the only one known to me in which, in three or four of
the species, there are what appear to be rudimentary staminodes at the base of
the inner perianth lobes, alternating with the filaments at their point of attach-
ment to the perianth tube. The staminodes of W. Pillansii are shown in fig. 3
on pi. XXII and in more detail in fig. <$a. In figs. 1 and la on the same plate
are shown the staminodes of W. marginata. In this species, in addition to the
structures which are presumed to be staminodes, there are small callus-like
projections below the filaments of the three stamens as well as below the
staminodes, those below the staminodes being smaller and more or less adnate
to the staminodes (see fig. ib).
In one or two species of Lapeyrousia (e.g. L. divaricata and L. anceps) there is a
small callus on each of the three lower perianth lobes and similar calli occur
in one or two species of Tritonia (e.g. T. securigera) . These calli should probably
be interpreted as ligular or stipular in origin. In many species of Moraea (also
in Ferraria and Homeria) there is a small shallow nectary at the base of the three
outer perianth segments, and in one or two species, such as M. tricuspidata (L. f.)
Lewis and M. confusa Lewis, there is a very small flap at the base of the outer
segments which is clearly ligular.
In almost all members of the Iridaceae the anther cells are parallel and slit
open from top to bottom to shed their pollen. Exceptions occur in Ferraria, in
which the anther-cells in some species are distinctly divaricated, and also in the
subgenus Dichone of Tritonia in which the cells open only by a small slit at the
base (see figs. 5 and $a on pi. XXII). In this as well as in other characters the
subgenus Dichone differs from the other subgenera of Tritonia, and in my opinion
it should be established as a separate genus. It is far more closely allied to Ixia
than to Tritonia.
In Babiana stricta and its varieties and allied species the anther-cells are widely
separated by the broad connective (see fig. 6 on pi. XXII). A characteristic
feature of the stamens in Exohebea is the apiculus, formed either by an extension
of the connective or, as in E. lata, by an extension of the anther-cells (see figs. 7
and ja on pi. XXII). An exception in this genus is E. parvifiora, in which, as
can be seen in figs. 8 and Sa, there is no apiculus.
Heterostyly has been recorded in two genera of the South African Iridaceae,
in Nivenia, and also in one species of Geissorrhiza, namely G. heterostyla L, Bol.
It is possible that it also occurs in Syringodea but it is necessary to examine more
material in this genus before any conclusion can be reached. In a few species of
Geissorrhiza, though the flowers are not heterostylous, the filaments in each
individual flower are of different lengths, two long and one considerably shorter.
This perhaps is a step towards heterostyly. The flowers are protandrous in three
or four genera, as for example Exohebea and Anapalina.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE IOI
One other important feature in connection with the stamens should be
mentioned. In some genera in the Iridoideae the filaments are completely
connate in a tube round the style, as in Homeria and Ferraria. In Moraea they
are more or less connate at the base. In Galaxia, in the Crocoideae, they are also
completely connate and in Ixia monadelpha in the Ixioideae they are connate in
the lower half. With the exception of the various cases mentioned above the
anthers or stamens do not present many characteristics which are of much value
for diagnosis.
6. The style and style branches
The structure of the style branches shows considerable variation and these
afford useful characters which have been largely used in systematic work for
the distinction of genera or small groups of genera. Many of the different forms
have been illustrated by Pax and Marloth and others who have worked on this
family, so it would be superfluous to discuss them in any detail here. On the
whole the structure of the style branches within the different genera is fairly
homogeneous so that this character is a fairly reliable one, although a few
exceptions have been recorded in some of the genera. In species in which these
exceptions occur a comparative examination of the vegetative organs will, as
a rule, remove any doubt as to the genus to which the plant belongs. In two or
three species of Lapeyrousia, for instance, such as L. Vaupeliana Dinter and
L. effurcata Lewis, the style branches are simple instead of bifid, yet the corm,
leaves, etc., are typical of the genus, or of the particular group of species within
the genus.
Two exceptions also occur in Geissorrhiza in which genus the style branches
are characteristically short, recurved, narrow and conduplicate; in G. Mathewsii
L. Bol. and G. Mathewsii var. eurystigma the style branches and stigmas are
broad, flat, thick, and crenate-edged. In Exohebea the stigmas are retuse (see
figs. 9 and ga on pi. XXII) but in most of the specimens of E. ramosa which have
been examined they were found to be shortly but distinctly bifid. The style
branches and stigmas of Moraea plumaria, shown in fig. 10 on pi. XXII, are
quite different from the crested style branches which characterize the genus.
A few other anomalies might be mentioned but those which have been
commented on in connection with the various organs of the flowers will serve
to confirm the statement made in the introduction to this paper, that it is often
impossible to describe any large group in this family, such as a tribe or genus,
without some proviso to cover the exceptions which occur. They also serve to
stress the necessity of taking into consideration the morphological characters of
the entire plant.
Section II. Systems of Classification
The different opinions held with regard to the classification of the Iridaceae
have already been mentioned in the introduction. To recapitulate, the three
main systems generally recognized are briefly as follows:
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE IO3
( i ) Bentham and Hooker's classification of the Iridaceae in Genera Plantarum
(1883). The family is divided into three tribes, Moraeeae, Sisyrinchieae and Ixieae.
The first two tribes are subdivided into several subtribes. This system is
followed by Baker in the Flora Capensis ( 1 896) and by Diels in the second edition
of Engler and Prantl's Pflanzenfamilien (1930), with only one or two minor
alterations.
(2) Pax, in the first edition of Engler and Prantl's Pflanzenfamilien (1889),
raised one of Bentham and Hooker's subtribes of the Sisyrinchieae to subfamily
status, Crocoideae, and combined the remaining subtribes of Sisyrinchieae with
those of Moraeeae, treating this also as a subfamily to which he gave the name
Iridoideae. This subfamily he subdivided into several tribes and subtribes. The
third group, corresponding with Bentham and Hooker's Ixieae, he also raised to
subfamily status, Ixioideae, and divided into three tribes. This system is followed
by Rendle in his Classification of Flowering Plants (1904) and by Marloth in - The
Flora of South Africa (191 5).
(3) Hutchinson, in his Families of Flowering Plants (1934), divided the family
into eleven tribes of equal status. Weimarck, in 1940, placed the shrubby genera
together in a separate tribe, Nivenieae, this bringing the number of tribes up to
twelve.
In attempting to work out a phylogenetic diagram to illustrate the natural
relationships of the South African members of the Iridaceae the various
morphological features discussed in this paper were all taken into consideration,
such as the type of inflorescence, the presence or absence of a pedicel, the nature
of the subterranean stem, the form of the leaf and bracts, and the structure of
the flower and its organs. Only the genera which occur in South Africa are
recorded in the diagram, which is given on plate XXIV. Although the position
of the majority of the genera which occur in other countries is fairly clear from
the descriptions, it is not easy to place all of them satisfactorily without having
examined them in person; since it has not been possible to do this I do not feel
justified in attempting to place them all in their correct positions.
The fairly numerous genera in South Africa include the most advanced as
well as some of the most primitive found in the family ; in addition there are
many genera which display characters intermediate between the latter and the
former and thus provide some evidence of the lines along which most of the
more highly evolved genera have developed. These South African genera are
representative of the whole family, and it is possible therefore to work out from
them the relationships of the genera and the groups to which they belong. It
was found that they fall fairly naturally into a number of groups, most of them
distinct and well defined.
By taking into consideration the correlated morphological characters in the
various groups they in turn can be divided into three major groups, two of them
large and consisting of three or more of the minor groups, and one small group
by itself which is morphologically quite distinct from the others. It is proposed
104 ANNALS OF THE SOUTH AFRICAN MUSEUM
to treat these three major divisions as tribes, Irideae, Nivenieae and Ixieae, and
the smaller groups in two of the tribes (i.e. Irideae and Ixieae) as subtribes. In
the diagram the boundaries of the tribes are indicated by double lines and those
of the subtribes by single lines.
On the whole, with some alterations, the arrangement arrived at approxi-
mates most closely to that of Pax, though Pax's subfamily Crocoideae has fallen
away and Weimarck's tribe Nivenieae is retained as distinct from the Irideae. Pax
combined the shrubby genera with Aristea, treating this group as one of the
tribes of the Iridoideae. The tribes Irideae and Ixieae correspond fairly closely with
Pax's subfamilies Iridoideae and Ixioideae, though with two important changes,
namely the removal of the shrubby genera from the former and the division of
the four genera of the Crocoideae between the two tribes.
Pax's subfamily Crocoideae presents a few problems and seems to be an artificial
group linking the Irideae and Ixieae together. The four genera placed in this
group, Galaxia, Syringodea, Romulea and Crocus, were separated from the Iridoideae
and Ixioideae respectively only on account of their abbreviated axis and reduced
types of inflorescences, but if all the morphological features of these plants are
taken into consideration it becomes apparent that the affinity of Galaxia is with
the Irideae and of the other three genera with the Ixieae, and that although
similar reductions have taken place in these plants it is very unlikely that they
have developed along the same direct line.
Galaxia has the same type of corm as Moraea and its allied genera, with corm
tunics similar to those of some species of Moraea. The dorsiventral leaves of
Galaxia, although reduced and very short, are also of the same type as those of
Hexaglottis, Moraea, etc. It differs from most other members of the Irideae only
in having undergone further reductions, resulting in the almost complete
suppression of the internodes of the axis and lateral branches, as well as a
reduction in the number of flowers in the cyme. Although the flowers have a
comparatively long perianth tube they are not sessile, for in all the specimens
examined a short pedicel was found, sometimes up to 5 mm. long, though more
often only 2 to 3 mm. long. Three species of the Irideae have been mentioned
in which there is distinct evidence that a similar reduction is taking place. In
Bobartia macrospatha and Moraea Cooperi the cymes are one-flowered, the pedicel
considerably reduced, and the flowers have a short perianth tube. In Hexa-
glottis virgata similar reductions have also occurred and the two-flowered cymes
are sessile in that species and the flowers have a short perianth tube. In Galaxia
the cymes are sessile and one-flowered, with the inner bract or spathe consider-
ably shorter than the outer one. ,The outer spathe resembles the leaf which
subtends the cyme and between the cyme and the axis is a pair of united
prophylls.
Another character of Galaxia which links this genus more closely to Homeria
and its allied genera is that the filaments are united around the style and the
anthers opposite the style arms, instead of alternating with them as they do in
Romulea and Syringodea. In Moraea, Homeria, Iris and allied genera (i.e. Bentham
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE IO5
and Hooker's tribe Moraeeae) the style arms are opposite the stamens, but in all
the other genera they alternate with them and are above the septa of the ovary.
Pax suggested that the second position is due to a twisting, the original position
as in Moraea, etc., being the normal, namely above the dorsal suture of the
carpels. This phenomenon requires further investigation, though it is very
probable that Pax's theory is correct. Bentham and Hooker attached a great
deal of importance to this character and it was their main basis for establishing
Moraeeae and Sisyrinchieae as separate tribes.
The morphological characters of Romulea and Syringodea differ in several
respects from those of Galaxia but are the same as, or very similar to, those found
in a few species of Hesperantha and Geissorrhiza in the Ixieae. The corms have been
described and their similarity to those of Geissorrhiza and Hesperantha discussed.
The structure of the leaves and the evidence they provide of an affinity with the
Ixieae were discussed by Arber. Although the inflorescence in Romulea and
Syringodea is not spicate as in nearly all the genera in the Ixieae, it has undergone
very extensive reductions, especially of some of the internodes of the main axis,
and the flowers are not arranged in cymes as in the Irideae. In Romulea the
comparatively long lateral branches, like the main axis, all terminate in a
solitary flower, which has a perianth tube but no pedicel, and is subtended by
a bract with what appears to be a pair of united bracteoles opposite, instead of
a single inner bract or spathe as in Galaxia. In some species of Romulea the bifid
or bicarinate nature of the bracteoles is fairly evident, in others not or barely
perceptible. In one or two species of Hesperantha and Geissorrhiza reduced
inflorescences very similar to those of Romulea occur, with the main axis and
lateral branches all terminating in a solitary flower. Still further reductions
have taken place in the inflorescence and entire plant in Syringodea, although
this genus possesses some characters which are regarded as being more primitive
than those of Romulea, such as the simple style branches (bifid in Romulea) and
the presence of a short stipe below the ovary. The latter probably represents
the final remnant of a pedicel, such as is found in Galaxia, but in Syringodea the
flowers are not in sessile one-flowered cymes as in Galaxia. Instead each flower
has a subtending bract with a pair of united bracteoles between the flower and
axis, but no outer and inner spathes as in Galaxia.
The rather brief analysis of the characters of Galaxia, Romulea and Syringodea
serves to illustrate the fact that Galaxia is morphologically distinct from the other
two genera and it is for this reason that it is proposed to abolish Pax's Crocoideae
and to place Galaxia in the Irideae, in the same subtribe as Homeria to which it
is probably most closely allied, and to place Romulea and Syringodea together in a
separate subtribe with the Ixieae. Crocus, the fourth genus of the Crocoideae, is
confined to the Northern hemisphere, where some species of Romulea also occur.
It is undoubtedly evolved from the same ancestral stock as Romulea, through
further reductions in the internodes of the aerial axis, accompanied by an
elongation of the perianth tube, as in Syringodea. As Arber remarked : 'The
extreme reduction of the whole Crocus plant and the length of the flower tube
106 ANNALS OF THE SOUTH AFRICAN MUSEUM
may be regarded as characters in which the genus has progressed further than
Romulea on the path of specialization.' Crocus should clearly be placed in the
subtribe of Ixieae with Romulea and Syringodea.
The main basis for separating the three tribes is the inflorescence, but there
are other correlated characters as well. In the Irideae the flowers are pedicellate
and arranged in cymes (rhipidia) with an outer and inner bract or spathe.
The perianth segments are usually free, though in a few of the genera there is
a perianth tube. In a few species in two or three of the genera and all species of
Galaxia the cymes are reduced to a single flower, this reduction being correlated
with a reduction in the length of the pedicel and the development of a perianth
tube. In Galaxia the perianth tube is long and the pedicel very short. In a few
species of Aristea the pedicels are extremely short (about i mm. long), though
throughout the genus the perianth segments are free. The plants are herbs,
with a rhizome and evergreen leaves, or a corm and deciduous aerial leaves.
The corm, in nearly all of the genera which have a corm, is formed through the
accumulation of food reserves in a single internode of an axillary bud which
develops at the same time as the aerial axis, the aerial axis dying away right
down to the base. This type of corm is associated with a dorsiventral
leaf.
In the Nivenieae the flowers are sessile, all have a perianth tube (very short in
Klattia), and are variously arranged in reduced compound inflorescences, not
in rhipidia as in the Irideae, nor in spikes as in nearly all of the Ixieae. The plants
are small perennial evergreen shrubs with a large basal, more or less subterra-
nean stem and woody aerial stems bearing numerous distichous imbricate leaves
of the simple isobilateral equitant type ; secondary thickening takes place in the
underground and aerial stems. The three shrubby South African genera in this
tribe form a fairly homogeneous systematic group with many morphological
features in common which are distinct from those of the other two tribes.
The flowers in the Ixieae are sessile, all have a perianth tube, and in nearly all
of the genera they are arranged in spikes, in almost every case with a pair of
united bracteoles between the flower and the axis. Exceptions occur in Pillansia
and Lapeyrousia in which there is evidence of a transition from a more complex
inflorescence to a spike. In Romulea and Syringodea, one or two species of
Hesperantha and Geissorrhiza, and also in the subgenus Sophronia of Lapeyrousia,
the whole inflorescence is considerably reduced and the aerial axis extremely
abbreviated. In Syringodea there is a short stipe or pedicel below the ovary.
In Exohebea, Anapalina and two allied genera the bracteoles are not derived from
a pair of united bracts or prophylls as in the other genera. All except one of the
genera in the Ixieae have corms, nearly all of which are formed through the
accumulation of food reserves in two or more abbreviated internodes of the basal
underground portion of the aerial axis. Pillansia and Dierama are evergreen
herbs but in all the other genera the aerial leaves are deciduous. The leaves
vary considerably in structure but all are derived from the isobilateral equitant
type. Dorsiventral leaves do not occur in the Ixieae.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE IO7
The genera in the Irideae are segregated into three subtribes, marked la, ib
and ic in the diagram (pi. XXIV). The confusion resulting from the misinter-
pretation of the tribe Sisyrinchieae was mentioned by Weimarck when discussing
the systematic position of Aristea in his monograph of that genus. He placed
both Aristea and Bobartia in the Sisyrinchieae and this arrangement is followed
here, except that this group is treated here as a subtribe, Sisyrinchinae {id). The
other two subtribes correspond more or less with two of Pax's subtribes of the
Iridoideae, namely Cipurinae (ib) and Iridinae (ic), the only change of any impor-
tance being the inclusion of G '-ataxia in the former. It is rather doubtful whether
there is sufficient justification for Pax's division of the tribe Tigridieae into two
subtribes (Tigridinae and Cipurinae), but this can only be decided when it is
possible to examine material of the various genera concerned.
The classification in the Irideae is based almost entirely on the form and
differentiation of the flowers. In the Sisyrinchinae the style branches are simple
and alternate with the stamens, while in the other two subtribes the style arms
are opposite the stamens and variously modified, i.e. bifid, crested, flattened,
etc. The Iridinae are separated from the Cipurnae on account of the more elaborate
style arms of the former, which are flattened, winged and petaloid, with con-
spicuous bifid crests and the stigmatic surface on a small projecting flap below
the crests. The largest genus in the Iridinae is Iris, which is widely distributed in
the Northern hemisphere; Moraea, closely allied to Iris, is well represented
throughout South Africa and slightly less so in Tropical and North Africa.
Pax divided the Ixioideae into three tribes, but Bentham and Hooker, Baker,
Diels, and one or two others, grouped all the genera together and treated them
as a single tribe. Hutchinson also segregated these genera into three tribes,
though his tribes are not the same as Pax's. One of them, Antholyzeae, was
established by Hutchinson for the most advanced genera of the Iridaceae. This
last arrangement will be referred to again shortly ; it is not followed here as it is
considered to be an artificial one, based entirely on the symmetry of the flowers.
As will be seen in the diagram (pi. XXIV), it is proposed to divide the Ixieae
into six subtribes. One of these, marked 3d, has been discussed already.
Excluding Galaxia this corresponds with Bentham and Hooker's tribe Croceae,
or Pax's subfamily Crocoideae, but since the conception of this subtribe is not
in accordance with the earlier ideas it would probably be better to rename it
and the name Romulineae is suggested as being the most suitable. Romulea is
a large genus and is well represented both in the Northern and Southern hemi-
spheres. The plants in this subtribe are all small and very much reduced, with
an extremely abbreviated axis (except in one or two South African species of
Romulea) , and the inflorescence is not spicate, the flowers being borne singly and
each arising between its subtending bract and a pair of united bracteoles.
The name Watsoniineae could be adopted for the subtribe 36 since this cor-
responds almost exactly with Pax's tribe Watsonieae. The inflorescence is
spicate, except in several species of Lapeyrousia, and the style branches bifid in
all the genera, the only exceptions being two or three species of Lapeyrousia.
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
In Lapeyrousia and Watsonia, especially the former, there is some evidence of the
inflorescence being derived from a more primitive ramified one. This has been
discussed in the section dealing with the inflorescence. The subgenus Anomatheca
has diverged so far from the other two subgenera of Lapeyrousia that it should
probably be treated as a separate genus, closely allied to Freesia. In the diagram
the range covered by Lapeyrousia is indicated by placing this genus in two
positions, linked together by a bracket. This same method is employed with
some of the other genera, such as Watsonia, Gladiolus, Babiana, etc., to indicate
that various progressive stages occur in these genera.
Fairly detailed descriptions of the subterranean stem and inflorescence of the
monotypic genus Pillansia have been given in this paper and it has been pointed
out that the subterranean stem provides some evidence that this plant is derived
from a perennial evergreen ancestral stock, either a herb with a rhizome or
possibly a small shrub with a thick undei ground stem such as occurs in members
of the tribe Nivenieae. In the inflorescence also there is evidence that a reduction
has taken place, the presence of vestigial traces of bracts suggesting an affinity
with Nivenia. It is considered that Pillansia represents an intermediate stage in
development between the tribes Nivenieae and Ixieae, but that it is nearer to the
latter and should be included in that tribe. It seems to be sufficiently distinct
from the other genera in the Ixieae to be placed in a separate subtribe by itself
(marked 3a in the diagram) , the most primitive in the Ixieae, but directly linked
with the tribe marked 36 through Watsonia. The only applicable name for this
subtribe is Pillansiineae.
A group of morphologically closely related genera, which is also regarded as
constituting a distinct subtribe, are the genera in the column marked $c
( Tritoniopsis, Exohebea and Anapalina) . This subtribe is characterized by having
a deep-seated corm, with tunics of coarse, wiry, fibrous strands, not reticulated
and extending up in a long neck, by the bracteoles being longer than the bracts,
and by the leaves which, in their external appearance as well as their internal
structure, are quite distinct from those of any other members of the Ixieae. The
structure and development of the basal leaves and the rudimentary cauline
leaves of some species of Anapalina and Exohebea have been discussed. In these
genera the bracts have developed adnate to a pair of prophylls and the bracteole
is not derived from a pair of united prophylls or bracteoles as in the other genera
in the Ixieae. It has been suggested that two species which N. E. Brown included
in the genus Chasmanthe (i.e. C. caffra and C. intermedia) should be transferred to
Anapalina as all the morphological features of these two species are the same as
those of Anapalina and, except for the similarity of the flowers, quite distinct from
those of Chasmanthe. By transferring these two species to Anapalina this genus
extends up into the ranks of the most advanced members of the Ixieae in which
the flowers are very irregular, with three or five of the perianth lobes much
smaller than the uppermost. The name Exohebineae is suggested as being the
most suitable for this subtribe.
There is little doubt that Brown's monotypic genus Tanaosolen should be
combined with Exohebea, the only difference between them being the fact that
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE IO9
the perianth tube is long in Tanaosolen and short in most species of Exohebea.
It has been pointed out that long- and short-tubed flowers occur in several
genera in the Ixieae and this character alone is not considered to be of much
importance. In most species of Exohebea the perianth tube is short but there are
two species with a long tube as well as one with a tube intermediate between the
long and short forms.
Another small group of genera with a combination of morphological features
in common which do not occur together in the other members of the Ixieae are
Babiana, Anaclanthe and Antholyza. They differ from the other genera in having
deep-seated corms with reticulate fibrous tunics (quite distinct from those of
Anapalina and Exohebea) and plicate leaves which are divided into two distinct
portions, a long, narrow sheathing base and an expanded pseudo-lamina. In
most species of Babiana the stem, leaves and bracts are pubescent; in Anaclanthe
and Antholyza only the stem and bases of the bracts are pubescent. Another
character peculiar to this group is the very short, compact and secund spike,
with the internodes between the flowers extremely reduced. This type of spike
occurs in Antholyza, Anaclanthe and the majority of species of Babiana. These
genera, which have clearly developed along the same line, the most advanced
being Antholyza, seem sufficiently distinct from the other genera in the Ixieae to
warrant being placed in a separate subtribe (marked 3/ in the diagram), to
which the name Babianiineae could be given, since this is by far the largest and
most widely distributed genus in the group.
Plicate leaves occur in one other genus in South Africa, in Curtonus in the
Ixieae, but the corm and corm tunics, bracts, inflorescence and flowers are not
the same as those of Babiana, Anaclanthe and Antholyza. According to Arber,
plicate leaves also occur in two or three genera in the Irideae which are not found
in South Africa. Arber commented on this occurrence of foliated leaves in the
two main groups of the family, which she considered to represent a case of
parallel development.
The subtribe of the Ixieae marked 30 in the diagram is a large one, but it was
found that any attempt to divide it into subtribes, as was done by Pax, and to a
further extent by Hutchinson, results in an artificial grouping of the genera
based entirely on the development or symmetry of the flowers. Pax divided this
group (which included Babiana and Antholyza, the latter a mixture of several
genera as was later pointed out by N. E. Brown) into two tribes, Ixieae and
Gladioleae, the division following the horizontal line in the diagram between the
groups of genera with actinomorphic or subactinomorphic flowers and those
with zygomorphic or subzygomorphic flowers. Hutchinson distributed the
genera of Pax's tribe Watsonieae between the tribes Ixieae and Gladioleae, though
he went a step further than Pax in establishing the tribe Antholyzeae in which he
placed all the most advanced genera which are shown in the uppermost section
in the diagram (i.e. from Anapalina to Antholyza), all of which have the three or
five lower perianth lobes considerably smaller than the uppermost.
Although the more primitive genera near the base of this group, i.e. Dierama,
Schizostylis, Hesperantha, Geissorrhiza and Ixia, are quite distinct from the advanced
I I O ANNALS OF THE SOUTH AFRICAN MUSEUM
genera at the top, various progressive stages in development can be traced in
this group where it is evident that many of the genera are closely linked together.
Dierama and Ixia are closely allied, as has been pointed out, and in Dierama there
is a transition from an evergreen plant with a perennial corm to a geophyte
with deciduous leaves and a corm which does not persist for more than a year.
Schizostylis and Hesperantha are also very closely allied, the only difference being
that the former has a rhizome and the latter a corm. The flat-based corms which
occur in some species of Hesperantha (as well as Romulea and Lapeyrousia) have
been discussed and it was pointed out that these give some indication of a transi-
tion from a rhizome to a corm. The borderline species between Geissorrhiza and
Gladiolus have also been mentioned and the fact that in some species of Geissor-
rhiza and Hesperantha the stamens and style are all directed to one side, which is
obviously a preliminary step towards the formation of a medianly zygomorphic
flower.
Pax included Babiana in the Gladioleae but in this genus there are species with
actinomorphic and others with zygomorphic flowers. In the genus Lapeyrousia
there are also species which fall into both categories. Pax's division also resulted
in the separation of Streptanthera from Sparaxis and Synnotia. On account of its
actinomorphic flowers Streptanthera was placed with the Ixieae and the other two
genera with the Gladioleae. These three genera are morphologically closely
allied, especially Streptanthera and Sparaxis which have almost identical corms,
corm tunics, leaves and bracts, and natural hybrids between these two genera
have been recorded in the National Botanic Gardens, Kirstenbosch.
It is evident that an attempt to divide the genera in the Ixieae into subtribes
based on the symmetry of the flowers results in a false division not only of
closely allied genera but also of species within one or two of the genera. The
same is true of the more advanced group of genera. It has been pointed out that
Gladiolus is extremely closely allied to Homoglossum and that Homoglossum passes
into Petamenes. In the section marked 3c, Anapalina passes upwards into the most
advanced region through the species at present known as Chasmanthe caffra and
C. intermedia. I consider therefore that the whole group of genera in the section
marked $e can only be regarded as a subtribe of the Ixieae, in which various
progressive stages occur as they do in the other subtribes. In three of them, 3c,
$e and 3/, the flowers have all developed along similar lines, resulting, in the
most advanced members, in medianly zygomorphic, very irregular flowers in
which three or five of the lower perianth lobes are reduced, sometimes consider-
ably so, while the uppermost is elongated and expanded. In each of these
respective groups or subtribes the other morphological features of the most
advanced genera are the same as those of the lower genera in the same group
and distinct from those of the other more advanced genera. For instance,
Antholyza and Anaclanthe (which are scarcely separable) are very closely con-
nected with Babiana but morphologically quite distinct from Anapalina, Homo-
glossum and Chasmanthe. It seems preferable, therefore, to place these most
advanced genera, as is shown in the diagram, at the top of the subtribes 3c, $e
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE I I I
and 3/. In my opinion this results in a more natural phylogenetic grouping of
the genera. The name Ixiineae is suggested for the subtribe 30, though the
conception of this subtribe is much broader than that of the tribe Ixieae of Pax
and Hutchinson and includes members of their tribe Gladioleae, as well as some
of the genera of Hutchinson's tribe Antholyzeae.
In the diagram one or two other slight alterations are proposed which will
necessitate some changes to the previous conception of one or two of the genera.
For instance, Anomalesia and Kentrosiphon are combined into one genus and the
subgenus Dichone is separated from Tritonia and raised to generic status, also the
subgenus Anomatheca of Lapeyrousia. N. E. Brown's genus Radinosiphon has been
included with a query as this requires further investigation. It is doubtful
whether it is sufficiently distinct from Gladiolus to warrant generic status.
With a few additions the scheme for classification which has been outlined
in the preceding pages could be made applicable to the whole family, though
this cannot properly be done without making an examination of some of the
genera which occur in other countries.
Summary
1 . In the prophylls of Witsenia maura what appears to be a rudimentary lamina
is apparent in the very early stages but it tends to disappear as the prophylls
develop.
2. From the development of the prophylls in Witsenia maura it appears that
the amplexicaul base of the petiole develops outside and adnate to an 'axillary'
ligule. In the shrubby genera, Witsenia, Nivenia and Klattia, an absciss layer is
formed at the base of the limb, below the sheathing portion.
3. It is suggested that the delay in the entry of the median vascular strand
into the leaf base is associated with the partial adnation of the axillary buds to
the axis. This adnation is evident in some of the shrubby genera, especially in
Nivenia Stokoei.
4. In the minute rudimentary cauline leaves of some species of Anapalina and
Exohebea there is further evidence of a laminar rudiment. In these plants the
bracts and rudimentary cauline leaves are adnate to the prophylls.
5. The basal leaves of Exohebea flexuosa, Anapalina triticea and A. Burchellii are
divided into a very slender solid petiole and an expanded pseudo-lamina.
6. The young dorsiventral leaves of Hexaglottis flexuosa (also other species of
Hexaglottis and several species of Moraea and Homeria) have a slightly swollen
area at the apex which is possibly a laminar rudiment. The apex of the leaf dies
off at an early stage.
7. Evidence is brought forward to substantiate Ingeborg Haeckel's theory
that the more primitive inflorescence in the Iridaceae is a much-branched one,
the spicate inflorescence of the Ixioideae being derived from this by various stages
of reduction.
112 ANNALS OF THE SOUTH AFRICAN MUSEUM
8. In some species of Babiana the prophylls and bracteoles are entirely free
and separate, providing some evidence that these are two distinct foliar
members which in most of the genera are more or less united owing to reduction
of the internodes.
9. Prophylls in various stages of fusion were found subtending cormlets in
Watsonia bulbifera. Where the fusion is only partial, two buds are present, but
a further overlapping and fusion results in the suppression of one of the buds.
10. Some vestigial traces of bracts in one or two species of Watsonia and in
Pillansia Templemanni furnish evidence that a reduction has taken place. The
inflorescence of Pillansia represents an intermediate stage between a compound
inflorescence like that of some species of Nivenia and the spicate inflorescence
of Watsonia.
1 1 . In some species of Lapeyrousia the branches and flowers, with their sub-
tending leaves or bracts, are more or less adnate to the axis. The inflorescence
is richly ramified in some species of Lapeyrousia and spicate in others, and there
are indications of a transition from the former type to the latter.
12. The fused bracteoles of Tulbaghia violacea are described briefly as they
provide further proof of fusion of adjacent overlapping bracts.
13. The reduction of internodes and the occurrence of short and long inter-
nodes in the Iridaceae are discussed.
1 4. The rhipidia and inflorescences of one or two species of Aristea, Bobartia
and Moraea are described. The apparently single rhipidium of the two latter
genera is probably derived by further reductions from 'binate rhipidia' such as
are common in Aristea.
15. The rhizome is the more primitive type of subterranean stem in the
Iridaceae and the corm a later development. The subterranean stem of Pillansia
represents a transition stage between a rhizome and corm.
16. The unusual 'corms' of Ferraria, which have no persistent scale leaves, are
described. These probably represent a transition stage from a rhizome to the
type of corm which occurs in some members of the Iridoideae.
17. In the Ixioideae the corm is formed through the accumulation of food
reserves in two or more abbreviated internodes in the underground basal part
of the aerial axis ; in the Iridoideae the food reserve is stored in a single internode
of an axillary bud.
18. The flat-based scale leaves of the corms of Lapeyrousia, Romulea and
Hesperantha, the spinous roots of Moraea ramosissima and the possible ligular
origin of the distinct inner layer of the scale leaf of Moraea plumaria are discussed.
19. Some anomalies observed in the flowers in some of the genera are
mentioned. In three genera in the Iridoideae previously described as having the
perianth segments free there are species (one in each genus) with a perianth
tube.
MORPHOLOGY, PHYLOGENY, TAXONOMY OF S.A. IRIDACEAE II3
20, The origin of the perianth tube and the probable derivation of either the
whole or part of it from the pedicel is discussed. A few controversial taxonomic
groups are mentioned.
21. A phylogenetic diagram is drawn to illustrate the natural relationships
of the South African genera of the Iridaceae and an amended system of classifica-
tion is proposed.
Bibliography
Adamson, R. S., 'Anatomy of Some Shrubby Iridaceae'. Trans. Roy. Soc. S. Afr., XIII, 175
(1926).
Arber, A., 'The Leaf Structure of the Iridaceae considered in relation to the Phyllode Theory'.
Ann. Bot., XXXV, 301 (1921).
, Monocotyledons. A Morphological Study. (1925.)
Baker, J. G., Flora Capensis, VI (Iridaceae) (1896).
Bentham, G., and Hooker, D. H., Gen. Plant., Ill, 681. (Classification of Iridaceae.) (1883.)
Bolus, L., i Pillansia TemplemannV ', Ann. Bolus Herb., I, 20 (191 5).
, l Moraea apetala', S.A. Gard. and Country Life, XIX, 385 (1929).
, 'Notes on "Aniholyza" (N. E. Brown's subdivision of the genus)', S.A. Gard. and Country Life,
XXIII, 46 (1933).
Bolus, L., and Mathews, W. F., 'Watsonia bulbifera', Ann. Bolus Herb., Ill, 140 (1923).
, 'Watsonia vivipara\ Ann. Bolus Herb., IV, 117 (1925-8).
Brown, N. E., 'Contributions to a Knowledge of the Transvaal Iridaceae'. Trans. Roy. Soc.
S. Afr., XX, 261 (1932).
, 'The genus Dierama\ Journ. Royal Hort. Soc, LIV, 1 (1929).
Diels, L. 'Classification of Iridaceae in Engler & Prantl Nat. Pflanzf.', ed. 2, 15a, 469 (1930).
Dinter, K., 'Lapeyrousia Vaupeliana\ Fedde Rep., XVIII, 436 (1922).
Foster, R. C, Revision of Geissorrhiza. Contr.from the Gray Herb. Harvard Univ. CXXXV, 1-78
(1940-
, Notes on Nomenclature of Iridaceae. Contr.from Gray Herb. Harvard Univ. CXI V, 38-41
(1936).
, Miscellaneous New Species and New Combinations. Contr. from Gray Herb. Harvard
Univ. CXXVII, 47-8 (1939).
Haeckel, 1., Flora, 125, 1-82 (1931). (Inflorescences of some of the Iridaceae.)
Hutchinson, J., Families of Flowering Plants, II (Monocotyledons) (1934).
Lewis, G. J., 'The genus Engy siphon and a Revision of the species placed under Acidanthera in
the Fl. Cap.' Journ. S.A. Bot., VII, 19 (1941).
, 'Notes on Bobartia lilacina and B. paniculata\ Journ. S.A. Bot., XI, 108 (1945).
, Iridaceae in Adamson and Salter's Flora of the Cape Peninsula (1950).
, 'A note on the rediscovery of Thunberg's "Gladiolus flexuosus' 1 ' 1 ( = Exohebea flexuosa) and a
description of the plant'. Ann. S. Afr. Mus., XL, 1 (1952).
Marloth, R., Flora of S. Africa, IV (Iridaceae) (1915).
Pax, F., Classification of Iridaceae in Engl. & Prantl 'Nat. Pfianzf, II, 5, 137 (1888).
Phillips, E. P., 'A note on N. E. Brown's sub-division of the genus Antholyza\ Bothalia, IV, 43
(1941).
Rendle, A. B., Classification of Flowering Plants, I (Iridaceae) (1904).
Scott, D. H., 'Notes on Spinous Roots'. Ann. Bot., XI, 327 (1897).
Scott, D. H., and Brebner, G., 'Secondary Tissue in certain Monocotyledons'. Ann. Bot., VII,
21 (1893).
Weimarck, H., 'Types of Inflorescences in Aristea and some Allied Genera'. Bot. Notiser, 616
(i939)- (Lund.)
, 'Monograph of the genus Aristea'. Lund. Univ. Arsskrft. N. F. Avd., 2, Bd. 36, Nr. 1 (1940).
, 'A Revision of the genus Nivenia Vent.'. Svensk. Botanisk. Tidskrift, Bd. 34, H. 4, 355 (1940).
The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irregular
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XXXIX.
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XL
PART III, containing: —
4. Iridaceae — New Species and Miscellaneous Notes. By G. J. Lewis.
(With five text-figures.)
ISSUED SEPTEMBER 1954
PRICE 5 j.
PRINTED FOR THE
TRUSTEES OF THE SOUTH AFRICAN MUSEUM
BY THE RUSTIC A PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE
4. Iridaceae — New Species and Miscellaneous Notes. By G. J. Lewis.
(With 5 text-figures)
Moraea macron yx Lewis sp. nov. (Iridaceae — Moraeeae)
M. macrochlamys affinis sed floribus majoribus et unguibus perianthii segmen-
torum multo longioribus differt.
Cormus globosus, 1-1-5 cm - diam., fere bulbillos emittens, tunicis extimis e
fibris paucibus pallidis crassis reticulatis compositis. Caulis brevissimus, omnino
foliis opertus; planta (cormo et flore incluso) 15-25 cm. alta. Vaginae basales 2,
pallidae membranaceae, obtusae, 1-2 et 3-5-6-5 cm. longae. Folia 2 caulina,
glabra, falcata, 12-25 cm - longa, tertia parte inferiore late vaginante, parte
superiore conduplicata, marginibus planis vel partim undulatis. Cyma unicum,
3-4 - florura, floribus sat grandibus, flavis immaculatis. Spatha exterior foliis
similis sed breviora dimidio parte inferiore vaginante; spatha interior recta,
saepe ventricosa, obtusa vel subacuta, 8-1 1 cm. longa, plerumque 1-1-5 cm -
diam., sed in plantis minoribus interdum 6 mm. diam. Perianthii segmenta
longiunguiculata, exteriora 4-5 cm. longa ungue 2-5-3 cm - longo, lamina ovata,
obtusa vel parum emarginata, 1-4-1-8 cm. lata; interiora 3-6-3-9 cm. longa,
lamina oblongo-ovata obtusa, 7-8 mm. lata. Filamenta 1-3-1-5 cm. longa, per
6-8 mm. coalita; antherae o-8-i cm. longae. Rami styli (cristis inclusis) 3-5-4
cm. longi, cristis acutis circa 2 cm. longis, 4-6 mm. latis. Ovarium 1-5 cm.
longum; pedicelli pallidi tenuissimi, 8-1 1 cm. longi.
Hab. Cape Province. Ceres Division: Cold Bokkeveld, near Elandsfontein,
about six miles north from top of Gydo Pass, in sandy ground, Lewis, South
African Museum No. 61920 {Type, in S.A.M. Herb.), 22 Sept. 1952; Cold
Bokkeveld, Leeurivier, Lewis, S.A.M. No. 57966, Sept. 1945; Cold Bokkeveld,
Schlechter 8890 (Kew) ; Cold Bokkeveld, top of Gydo Pass, Leipoldt 4082 (in
Bolus Herb.), Sept.; Cold Bokkeveld, near Elands Kloof, Leighton 1287 (Bolus
Herb.), Sept.
Corm globose, 1-1-5 cm - diam., with the outer tunics composed of a few pale,
coarse, reticulate fibres; cormlets usually present. Stem simple, very short,
enclosed within the leaf sheaths; height (including corm and flower) 15-25 cm.
Basal sheaths 2, membranous, colourless, obtuse, 1-2 and 3*5-6-5 cm. long.
Leaves 2, cauline, glabrous, falcate, 12-25 cm - l° n g> tne lower third wide and
sheathing, conduplicate above, the margins smooth or partly undulate ; lowest
leaf sometimes with an axillary cormlet. Cyme solitary, 3-4 - flowered, the
flowers fairly large, yellow, without markings. Outer spathe like the leaves but
slightly shorter, with the lower half ventricose ; inner spathe erect, often ventri-
cose, obtuse or subacute, 8-1 1 cm. long, usually 1-1-5 cm - diam., in smaller
plants sometimes only 6 mm. diam. Perianth segments with long claws, the outer
"5
VOL. XL. PART 'I
n6
ANNALS OF THE SOUTH AFRICAN MUSEUM
Figure i. Moraea macronyx Lewis, i. Plant in flower. 2. Outer perianth segment. 3. Inner
perianth segment. 4. Flower with perianth segments removed, showing pedicel, ovary, stamens
and style branches. 5. Ovary and part of pedicel of faded flower, showing slight contractions
in the pedicel. All drawings natural size. Del. G. J. Lewis-
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES I I J
ones 4-5 cm. long with the claw 2-5-3 cm - l° n g an d an ovate obtuse, or slightly
emarginate blade 1-4-1-8 cm. wide; inner segments 3-6-3-9 cm. long, the claw
widening gradually into an oblong-ovate blade 7-8 mm. wide. Filaments
1-3-1-5 cm. long, united for half their length; anthers o-8-i cm. long. Style
branches (with crests) 3-5-4 cm. long, the crests acute, about 2 cm. long, 4-6 mm.
wide. Ovary 1-5 cm. long; pedicels colourless, very slender, 8-1 1 cm. long.
This species differs from M. macrochlamys Baker, to which it is most closely
allied, in having larger flowers and very much longer claws to the perianth
segments. The claw of the inner segments widens gradually into the blade, but
in the outer ones the long, narrow claw passes abruptly into the broad, ovate
blade. The mature flower, with its ovary, is well exserted from the spathes but
as the flower fades the long, slender pedicel begins to contract and the ovary
is withdrawn into the spathes, the capsule developing and ripening within the
ventricose sheath of the inner spathe. When the flower is mature the pedicel
has smooth margins but slight undulations and wrinkles have been observed in
the pedicels of flowers which have faded. This phenomenon of a contractile
pedicel is not common to the whole genus but it is almost certain that it occurs
also in the allied species M. macrochlamys and M. ciliata, as in herbarium speci-
mens examined the only capsules found were enclosed within the spathes, near
the base.
Ferraria foliosa Lewis sp. nov. (Iridaceae — Moraeeae)
F. undulatae L. affinis sed sine foliis basalibus et foliis caulinis valde patentibus
differt.
Cormus globosus, 2-5-3 cm - diam. Caalis circa 40 cm. altus, foliis caulinis
imbricatis opertus, spiraliter tortus, prope apicem aliquot ramis brevibus
instructus. Vaginae basales 2, submembranaceae, 6 et 1 1 cm. longae. Folia
14-20, omnia caulina, glauca, 17-6 cm. longa, obtusa et mucronulata vel
subacuta, tertia parte inferna vaginante ad 5 cm. lata marginibus pallidis
membranaceis, parte superiore valde patente, ad 6 mm. lata. Cymata 2 -flora.
Spatha exterior 2-8-3 cm. longa, 2 cm. lata apice acuta leviter incurva; spatha
interior 4*8-5 cm. longa, ad 3 cm. lata. Perianthii segmenta purpureo-rubida
marginibus fulvis, foetida, exteriora 3*3-3-8 cm. longa, supra unguem nectari-
ferum 1-1-4 cm. lata, interiora 2-8-3-3 cm - longa, o-8-i cm. lata. Filamenta
8-9 mm. longa, per 7 mm. coalita ; antherae suborbiculares, circa 2 mm. longae,
lobis parallelis. Rami styli 4-5 mm. longi cristis inclusis. Ovarium 1-6-1-8 cm.
longum, 3-3-5 mm. diam.; pedicelli 2 cm. longi.
Hab. Cape Province. Piketberg Division: Elands Bay, near Verloren Vlei,
in sandy ground, Lewis, South African Museum No. 60869 ( TyP e i m S.A.M. Herb.),
Sept. 1951 ; north-east side of Verloren Vlei at Matjesgoed Drift, Pillans 8127
(in Bolus Herb.), Oct.
Corm globose, 2-5-3 cm - diam., attached to 2 or 3 older corms and growing
more or less horizontally. Stem about 40 cm. high, spirally twisted, entirely
enclosed within the wide imbricating sheaths of the cauline leaves, with several
n8
ANNALS OF THE SOUTH AFRICAN MUSEUM
Figure 2. Ferrariafoliosa Lewis. 1. Plant — half natural size. 2. Flower — natural size. 3. Outer
perianth segment — natural size. 4. Flower with perianth segments removed, showing pedicel,
ovary, stamens and style crests — natural size. 5, Stamens and style crests — X i£. 6. Style arms
and crests, from above, X 2. Del. G. J. Lewis.
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES Iig
short flowering branches at the top. Basal sheaths 2, more or less membranous,
6 and n cm. long. Leaves 14-20, all cauline, glaucous, 17-6 cm. long, obtuse
and mucronulate or subacute, the lower third forming an inflated sheath, up to
5 cm. wide, with white membranous margins, the upper flat part spreading
more or less horizontally, 6 mm. wide. Cymes several, 2-flowered. Outer spathe
2-8-3 cm. long, 2 cm. wide, the apex acute, slightly incurved; inner spathe
4*8-5 cm. long, up to 3 cm. wide, more inflated than the outer. Flowers dark
maroon with yellow-brown margins, unpleasantly scented, with the nectari-
ferous claws of the segments curved up to form a cup about 1-5 cm. diam. ;
outer segments 3 # 3-3*8 cm. long, 1-1-4 cm - wide just above the claw; inner
segments 2-8-3-3 cm. long, o-8-i cm. wide. Filaments 8-9 mm. long, united for
7 mm.; anthers suborbicular, about 2 mm. long, with parallel cells. Style arms
4-5 mm. long, including the fimbriate crests. Ovary 1 -6-i -8 cm. long, 3-3-5 mm.
diam. ; pedicels 2 cm. long.
F. foliosa is allied to F. undulata, a species fairly common along the south-
western coast from the Malmesbury Division to Danger Point in the Galedon
Division, but differs in having no basal leaves and the cauline leaves spreading
out more or less horizontally above their ventricose bases. Owing to a slight
twist in the stem, the distichous leaves are spirally arranged, an arrangement
which is not easy to represent in a drawing and which is not apparent in dried
herbarium specimens. Corms were collected during July 1950, when the plants
were not fully developed, and grown at the South African Museum Herbarium,
where they flowered from the end of August to September 1951, and again the
following year. These plants are probably smaller than they would be if grown
in their natural habitat; a plant with fruits in the Bolus Herbarium, collected
near the type locality during October, is considerably taller and more robust
(more than 1 metre high) with correspondingly longer leaves and
branches.
Its natural habitat is in coastal sand, and the same species was observed at
two or three localities along the coast from Elands Bay to Lamberts Bay in
the Clanwilliam Division.
Ferraria brevifolia Lewis sp. nov. (Iridaceae — Moraeeae)
Cormus depresso-globosus, 1-5-2 cm. diam. Caulis brevissimus omnino foliis
opertus; planta (cormo et inflorescentia incluso) 12-19 cm. alta. Vaginae
basales 2 vel 3, pallidae membranaceae, acutae, 2-5-8-5 cm. longae. Folia plura
omnia caulina brevissima, patentia, 2-6 cm. longa laminis 1-1-5 cm - longis ad
1 cm. latis oblique ovatis marginibus crassis et apicem versus acutis leviter
incurvis. Cymata pauca, 2-flora. Spatha exterior foliis similis; spatha interior
5-5-5 cm. longa, obtusa vel acuta marginibus pallidis membranaceis sed apicem
versus crassis. Perianthii segmenta acuminata, flava, marginibus aurantiacis;
exteriora basi viride maculata, 3*3-3-7 cm. longa ungue circa 1-3 cm. longo,
lamina supra unguem ad 1 cm. lata; interiora immaculata, circa 3-3 cm. longa,
4-5 mm. lata. Filamenta 1 cm. longa, omnino coalita; antherae aurantiacae,
20
ANNALS OF THE SOUTH AFRICAN MUSEUM
Figure 3. Ferraria brevifolia Lewis. 1. Plant. 2. Flower, from above. 3. Outer perianth segment.
4. Inner perianth segment. 5. Stamens and style crests, X 2. 6. Anther cells, x 2. 7. Gynaeceum
and pedicel. 8. Style arms and crests, x 2. All drawings natural size except where stated.
Del. G. J. Lewis.
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES 121
divaricatae. Ovarium 2 cm. longum, apicem versus in rostro sterili 8-9 mm.
longo productum; pedicelli 2-5-3 cm - l° n gi-
Hab. Namaqualand: 2 miles south of Nieuwerust,* Lewis, South Afr. Mus.
No. 57943 (Type, in S.A.M. Herb.), 7 Sept. 1945; between Bitterfontein and
Garies, Lewis, N.B.G. No. 2065/32 (in Bolus Herb.), Sept. 1932; Bitterfontein,
Schlechter 11 031 (Kew and Bolus Herb.).
Corm depressed-globose, 1-5-2 cm. diam. Stem very short, hidden by the
leaves; height (including corm and inflorescence) 12—19 cm. Basal sheaths 2
or 3, colourless, membranous, acute, 2-5-8-5 cm. long. Leaves several, all short
and cauline, the lower ones spreading horizontally, the upper suberect, 2-6 cm.
long, the blades 1-1-5 cm - l° n g an d up to 1 cm. wide, obliquely ovate with
slightly incurved acute tips and thickened margins. Cymes few, 2 -flowered.
Outer spathe like the leaves; inner spathe 5-5-5 cm. long, obtuse or acute, with the
margins membranous and colourless but thickened towards the apex. Perianth
segments acuminate, yellow with orange margins; outer segments with a few
small green spots on the claw and base of the blade, 3-3-3-7 cm. long, the claw
about 1-3 cm. long, the blade above the claw up to 1 cm. wide; inner segments
not spotted, about 3-3 cm. long, 4-5 mm. wide. Filaments 1 cm. long, com-
pletely united; anthers bright orange, divaricate. Ovary enclosed within the
spathes, 2 cm. long, with a sterile beak-like prolongation 8-9 mm. long above;
pedicels 2-5-3 cm - long.
A very distinct species readily recognized by its short spreading leaves with
very short obliquely ovate blades which have conspicuous cartilaginous margins.
The attractive yellow and orange flowers have a fairly pleasant scent.
Ixia leipoldtii Lewis sp. nov. (Iridaceae — Ixieae)
Cormus globosus, 1-1-4 cm « diam., tunicis e fibris numerosis sat tenuibus
reticulatis compositis. Caulis 1 1-25 cm. altus, simplex vel ramo brevissimo
suberecto, interdum 2 ramis. Vaginae basales 1 vel 2, 1-5-4-5 cm. longae.
Folia 3, mediocriter robusta, 7-20 cm. longa, 3-5 mm. lata, infimum sub-
falcatum, secundum suberectum, supremum erectum fere vel omnino vaginans.
Spica 2-4 - flora. Bractea membranacea, fulva vel fulvo-lilacina, 0-7-1 cm.
longa, conspicue 3-nervata, acuta vel obtusa vel truncata, plerumque 3-cuspidata
sed interdum cuspide unica induta; bracteolae bracteae similes sed 2-nervatae
et breviter bicuspidatae. Perianthii tubus flavus, 1-1-2 cm. longus basi filiformis
superne infundibuliformis ; segmenta aequalia, alba, tertia parte inferna rubra,
orbicularia vel suborbicularia, 1-5-1-7 cm. longa, ad 1-4 cm. lata, basi ungue
brevissimo et lato. Stamina brevia, antherarum basibus in tubo perianthii
inclusis; filamenta 3-4 mm. longa; antherae 3-4 mm. longae. Stylus apicem fila-
mentorum attingens ramis circa 2 mm. longis. Ovarium 3 mm. longum, 2 mm.
diam.
* Now spelt Nuwerus.
122 ANNALS OF THE SOUTH AFRICAN MUSEUM
Hab. Cape Province. Prince Albert Division: Eikerkraal, between Prince
Albert and Klaarstroom, Leipoldt, Bolus Herb. No. 21808 {Type, in Bolus Herb.),
27 Sept. 1935.
Corm globose, 1-1-4 cm - diam., with numerous tunics of fairly fine reticulate
fibres. Stem simple or with 1 or sometimes 2 very short suberect branches,
11-25 cm. high. Basal sheaths 1 or 2, 1*5-4*5 cm. long. Leaves 3, fairly firm,
7-20 cm. long, 3-5 mm. wide, the lowest subfalcate, the second suberect, the
third erect and entirely or almost entirely sheathing. Spike 2-4 - flowered.
Bract membranous, brownish or brownish purple, 0-7-1 cm. long, conspicuously
3-veined, acute, obtuse or truncate, usually shortly 3-cuspidate but sometimes
with a single cusp ; bracteoles like the bracts but 2 -veined and shortly bicuspidate.
Perianth tube yellow, 1-1-2 cm. long, filiform at the base, the upper half funnel-
shaped; lobes equal, white with the lower third claret-coloured, orbicular or
suborbicular, 1-5-1-7 cm. long, up to 1 -4 cm. wide, with a very short broad claw.
Stamens short, with the basal quarter or third of the anthers not exserted from
the perianth tube; filaments 3-4 mm. long; anthers 3-4 mm. long. Style reaching
to the top of the filaments with branches about 2 mm. long. Ovary 3 mm. long,
2 mm. diam.
Although a bright, contrasting patch of colour at the base of the perianth
lobes is a fairly common feature in the section Euixia, it has not hitherto been
recorded in the section Morphixia, to which this species, with its distinctly
funnel-shaped perianth tube, belongs. The lobes are white, with a conspicuous
claret-coloured patch at the base, and short coloured veins extending upwards
into the base of the white part. Another character which distinguishes this
species from all others in the section is the width of the perianth lobes, the broad,
almost orbicular, lobes presenting a striking contrast to the more or less oblong
lobes of the other species in the group. This attractive and apparently rare
species has been collected only once and is named in honour of the collector,
the late Dr. G. L. Leipoldt, who will always be remembered in this country as
a writer and poet and a keen observer and lover of nature.
Thanks are due to the Curator of the Bolus Herbarium, Dr. L. Bolus, for the
loan of the type material and for the notes made by her on the colour and shape
of the living flowers.
Gladiolus symmetranthus Lewis sp. nov. (Iridaceae — Ixiae)
G. erectifloro et G. trichonemifolio affinis sed floribus omnino aequilateralibus
flavis, in faucibus purpureis, differt.
Cormus globosus vel subglobosus, 1-1-5 cm - diam. Caulis gracilis, 16-26 cm.
altus. Vaginae basales 2, obtusae, 2-3 cm. et 3-6 cm. longae. Folia 2 vel 3,
infimum dimidiam partem caulis vaginans, lamina subtereti valde striata, quam
caule duplo longiore; folia cetera multo breviora laminis brevibus subulatis
spicam non superantibus. Spica 1-3 - flora, floribus aequilateralibus flavis, in
faucibus purpureis. Bractea mediocriter rigida, crebre nervosa, acuta, 2-5-3 cm -
longa, 7-8 mm. lata; bracteolae parum angustiores obscure 2-dentatae. Perianthii
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES
123
tubus rectus, infundibuliformis, circa 1-5 cm. longus; segmenta subaequalia
oblonga obtusa vel interdum leviter emarginata, 2-5-3 cm - longa, 1*1—1 -3 cm.
lata. Stamina recta, aequilateralia, filamentis 8-9 mm. longis antherisque
8-9 mm. longis. Stylus rectus, prope apicem antherarum attingens ramis
3-4 mm. longis ; ovarium 4 mm. longum.
Hab. Cape Province. Stellenbosch Division: near Koelenhof, between
Stellenbosch and Mulders Vlei, in marshy ground on flats, Lewis, South Afr.
Mus. No. 60700 (Type, in S.A.M. Herb.), 14 Sept. 1950; Mulders Vlei, in
somewhat marshy ground, Duthie 940 (Bolus Herb.), Sept.; near Koelenhof,
Figure 4. Gladiolus symmetranthus Lewis. 1. Corm and base of stem. 2. Bract. 3. Bracteoles.
4. Flower, laid open. 5, Flower, side view, with upper part of the perianth lobes removed to
show the stamens and style, X 2. All drawings natural size except where stated. Del. G.J. Lewis.
Leighton, s. n. (Bolus Herb.), Sept.; Elsenburg, Grant 2512 (Bolus Herb.), Sept.;
between Paarlberg and Paardeberg, Drege 8457 (Geneva), Aug.— Sept.
Corm globose or subglobose, 1-1*5 cm - diam.; tunics coarse, usually con-
centric above with the lower half toothed. Stem slender, simple, 16-26 cm. high.
Basal sheaths 2, obtuse, 2-3 and 3-6 cm. long. Leaves 2 or 3, the lowest sheathing
the lower half of the stem, with a strongly ribbed subterete blade, more than
twice as long as the stem, the upper 1 or 2 leaves very much shorter with
subulate acuminate blades not reaching above the inflorescence. Spike
1-3 - flowered; flowers actinomorphic, bright yellow with the throat dark
purple-maroon. Bract firm, closely veined, acute, 2-5-3 cm - l° n g> 7~8 mm.
wide; bracteoles similar but slightly narrower, obscurely bifid. Perianth tube
straight, funnel-shaped, about 1-5 cm. long; lobes subequal, oblong, obtuse or
sometimes slightly emarginate, 2-5-3 cm - l° n g> i*i-i*3 cm. wide. Stamens erect,
symmetrically arranged, reaching to the middle of the lobes ; filaments 8-9 mm.
124 ANNALS OF THE SOUTH AFRICAN MUSEUM
long ; anthers 8-9 mm. long. Style erect, reaching nearly to the top of the anthers ;
style branches alternating with the anthers, 3-4 mm. long; ovary 4 mm. long.
This species is closely allied to G. erectiflorus Baker and G. trichonemifolius Ker,
and G. linearis (L. f.) N. E. Br. It differs from the two former in having actino-
morphic flowers, as well as in the colour of the flowers, those of G. erectiflorus
and G. trichonemifolius being yellow without a purple throat. In these two species
the perianth tube is usually somewhat longer than in G. symmetranthus and the
flowers are slightly zygomorphic with the three upper perianth lobes a little
broader than the lower ones. The third species previously described in this
small and very distinct group is G. linearis (L. f.) N. E. Br. (G. biflorus Klatt),
and in this species, as in G. symmetranthus, the flowers are actinomorphic, though
they are considerably smaller and of a different colour, either pale or bright
lilac. In the Flora Capensis (vi, p. 146), Baker remarked that this species was on
the edge of the genus in the direction of Geissorrhiza. There is undoubtedly a
close affinity between these four species and all of them have the same habitat,
in marshy ground on flats, either in or on the margins of shallow depressions
which are filled with water during the late winter and spring months.
Gladiolus jonqjjilodorus Eckl. ex Lewis sp. nov. (Iridaceae — Ixieae)
G. Pillansii proxime accedit sed foliis radicalibus 2, bracteis leviter brevioribus,
perianthii segmento supremo obtuso, et perianthii colore, distinguitur.
Cormus globosus, 1-5-2 cm. diam., tunicis e fibris pallide brunneis papyraceo-
membranaceis compositis. Caulis gracilis, 28-65 cm. altus. Vaginae basales 2,
obtusae, 6-5-13 cm. longae. Folia radicalia plerumque 2, glabra, teretia,
4-canaliculata, ad 60 cm. longa, 1-5-2 mm. diam., post anthesin evoluta;
folia caulina 3, omnino vaginantia, acuta, supremum 2-5-8 cm. longum, 2
inferiora longiora. Spica secunda, 5-9 - flora, floribus odoratis pallide flavis
vel gilvis, segmentis inferioribus citrinis maculatis. Bractea acuta, 1-2 cm.
longa, 6-7 mm. lata; bracteolae similes sed leviter breviores. Perianthii tubus
curvatus superne infundibuliformis, o-8-i cm. longus; segmenta 3 superiora
obovata, supremum obtusum, 2-2-5 cm. longum, 1-2-1-4 cm. latum, cetera
acuta vel subacuta; 3 inferiora 1-4-1-8 cm. longa, 6-8 mm. lata, acuta,
recurva, unguiculata, unguibus per 5 mm. connatis. Stamina arcuata filamentis
circa 1-3 cm. longis antherisque 6-8 mm. longis. Stylus apicem antherarum
attingens, ramis 2 mm. longis, stigmata obcordata.
Hab. Gape Province. Cape Division: Gape Flats, in damp places, £eyher 450
(Type, in S.A.M. Herb.); near Zeekoe Vlei, Pappe (G. fragrans MS) (S.A.M.
No. 21352) Feb.; Zeekoe Vlei, among Restionaceae, Purcell (S.A.M. No.
62030), 27 Dec. 191 7; south of Koeberg, on sandy flats, Pillans, s. n. (Bolus
Herb.); near Vygeskraal Farm, W. Dod 2392 (Bolus Herb.); Guthrie 1386
(Bolus Herb.); Pillans (Bolus Herb. No. 107 13); flats at Kitchings, Bergvliet,
Purcell (S.A.M. No. 62029), 27 Dec. 1915; dry, sandy ground near Schusters-
kraal, Scarborough, Minicki (S.A.M. No. 60912), 12 Jan. 1951; near roadside,
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES 1 25
i mile east of Kommetjie, among reeds in sandy ground, Linley (S.A.M. No.
62061), 10 Jan. 1953; near Potsdam, Mellish (S.A.M. No. 62060), 30 Dec. 1952.
Corm globose, 1*5-2 cm. diam., with thin, membranous, pale-brown tunics.
Stem simple, slender, 28-56 cm. high, up to 3 mm. diam. at the base. Basal
sheaths 2, obtuse, 6*5-13 cm. long. Basal leaves usually 2, glabrous, terete,
4-grooved, up to 60 cm. long, 1-5-2 mm. diam., appearing after the flowering
season and if present with the flowers then brown and dead; cauline leaves 3,
completely sheathing, the uppermost 2-5-8 cm. long, the others longer. Spike
secund, 5-9 - flowered. Bract acute, 1-2 cm. long, 6-7 mm. wide; bracteoles
similar, slightly shorter. Flowers very sweetly scented, cream-coloured or pale
yellow, sometimes flushed with pink or mauve outside, the lower lobes with
bright yellow transverse bands near the middle. Perianth tube curved, o-8-i cm.
long, the upper half funnel-shaped ; 3 upper perianth lobes obovate, the two
side lobes 1 -8-2 cm. long, 1 cm. wide, acute or subacute, the uppermost 2-2-5 cm.
long, 1-2-1-4 cm. wide near the apex; 3 lower lobes acute, recurved, ungui-
culate with the claws united for about 5 mm. at the base, 1-4—1-8 cm. long,
6-8 mm. wide. Stamens arcuate; filaments about 1 -3 cm. long; anthers 6-8 mm.
long. Style reaching to the top of the anthers, with branches 2 mm. long and
obcordate stigmas. Ovary fusiform, 5 mm. long, 1-5-2 mm. diam.
Although Ecklon distributed this species under the name jonquilodorus he did
not describe it, nor has any description been published since it was first collected
more than a hundred years ago. In the Flora Capensis (vi, p. 144 (1896) ) his
MS. name is cited under Gladiolus brevifolius J acq., as well as Pappe's MS. name
for the same plant, G.fragrans. Ecklon's name is now given to this species which
is quite distinct from G. brevifolius, from which it differs in having very much
thinner, more or less membranous corm tunics, two terete basal leaves instead
of a single, flat, narrow leaf, and also in the character of the sweet-scented
yellowish or cream-coloured flowers.
It is very much more closely allied to G. Pillansii Lewis, which has the same
habitat, i.e. dry sandy soil among Restionaceae, and occurs in some of the same
localities, but with a later flowering period ; G. jonquilodorus flowers from the
end of December to the middle of February and G. Pillansii from March to
April. Apart from the difference in the flowering period and the colour of the
flowers, those of G. Pillansii being blue with yellow bands on the three lower lobes,
G. jonquilodorus has two basal leaves instead of one, the flowers more closely
arranged in the spike, with slightly shorter bracts and bracteoles, and the upper-
most perianth lobe widest at the apex and obtuse instead of acute and widest
shortly below the apex. A pink-flowered variety of G. Pillansii has recently been
described and figured (G. Pillansii var. roseus Lewis, Fl. Plants of Afr., vol. 29,
p. 1 1 59). This was found in 1951 in the same locality as G. jonquilodorus, near
Schusterskraal in the southern part of the Cape Peninsula, but it flowers a
month later and has a different habitat, being confined to damp ground near
the vlei. In the Flora of the Cape Peninsula, published in 1 950, neither of these
species was mentioned as up to that date there was no record of G. Pillansii var.
126 ANNALS OF THE SOUTH AFRICAN MUSEUM
roseus, and it seemed extremely doubtful whether G.jonquilodorus occurred within
the area, all the early records except one being from localities just outside the
area. However, in January 1951 and 1952 it was collected by Miss J. Minicki
near Schusterskraal, and it was collected again in January 1953, near Kommetjie.
Gladiolus brevifolius Jacq. var. robustus Lewis var. nov.
A typo tota planta robustiore caule spicaque longiore, spica sat dense 12-20 -
flora, segmento perianthii supremo perspicue cucullato, obtuso, quam infimum
fere duplo latiore, distinguitur.
Hob. Cape Province. Malmesbury Division: near Saldanha Bay, Pole Evans
{Type, in Bolus Herb. No. 21465, and S.A.M. Herb. No. 51344), 7 Jan. 1935;
F. Bolus (Bolus Herb. No. 21465); in damp places near the salt-pan at Yser-
fontein, Thompson, s. n. (Bolus Herb.), Jan. ; Paarl Division: between Durban-
ville and Paarl, Lewis, s. n. (Bolus Herb.), March; Galedon Division: Elgin
Forest Reserve, Lewis (S.A.M. Herb. No. 60885), May.
Corm subglobose, 3 cm. diam. ; tunics reddish-brown, thin, splitting into
fibrous strands in the lower half. Stem 80-85 cm - high (including spike), 3—4 mm.
diam. at the base. Basal sheaths 2 or occasionally 3, the two lower obtuse,
2-2-5 anc * 4-8 cm. long, the third when present 16-19 cm - l° n §- acute, brown.
Basal /^produced after the flowering period, about 40 cm. long, 7-9 mm. wide,
spirally twisted towards the apex, shortly and sparsely pilose, usually with 4
fairly prominent veins, sometimes up to 9-veined. Cauline leaves 2 or 3, 13-2 cm.
long, closely sheathing, the lower one or two acute, the uppermost with a short,
acuminate blade. Spike secund, fairly densely 12-20 - flowered. Bract 2-1-4 cm.
long, 6 mm. wide, subacute; bracteoles 1-5-1-3 cm. long, 5-6 mm. wide, com-
pletely united. Flowers pink, the 3 lower lobes marked with yellow and crimson
streaks. Perianth tube 1 cm. long, curved just below the throat; uppermost lobe
obovate, hooded, 2-7 cm. long, i-6 cm. wide; upper side lobes subacute, 2 cm.
long, i-2 cm. wide; 3 lower lobes 1-5-1-7 cm. long, 7-9 mm. wide, with acute
or subacute recurved tips. Filaments 1-4 cm. long; anthers 7 mm. long. Style
branches 3 mm. long, just overtopping the anthers.
G. brevifolius Jacq. is the best known of the Cape autumn-flowering species
and was grown in England and Europe more than 150 years ago and figured
in three or four of the early botanical works. It is still fairly common on
the Cape Peninsula, especially in the southern area, and in various localities
in the Caledon Division. The flowers, which have little or no scent, vary
from pale to deep pink, usually with yellow markings and crimson streaks
on the lower lobes. The variety differs from the typical in being a considerably
taller and more robust plant, with a longer spike which has from twelve to
twenty flowers, whereas in the typical there are rarely more than twelve flowers
in the spike, the usual number being between six and twelve. The flowers them-
selves are very similar but in the variety the uppermost perianth lobe is slightly
more rounded and hooded, and almost twice as wide as the lowest lobe.
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES 127
The flowering season for G. brevifolius is from March to May, but plants have
occasionally been found in flower between June and August, in which case the
solitary basal leaf is sometimes present with the flowers. The flowering season
for the variety is from January to March but, as in the typical, there are occa-
sionally a few plants which flower out of the usual season, and there is one record
of the variety flowering in May. The solitary narrow basal leaf, which appears
after the flowering period, is variable in the variety, both as regards the number
of the rather prominent veins (from 4 to 9) and the pubescence, some leaves
being almost glabrous.
Gladiolus meridionalis Lewis sp. nov. (Iridaceae — Ixieae)
G. maculato Sweet proxime accedit sed floribus majoribus, segmentis latioribus,
perianthii tubo subcylindrico et perianthii colore, distinguitur.
Cormus globosus, 1—1*5 cm - diam., tunicis e fibris brunneis tenuibus reticulatis
compositis. Caulis gracilis, 30-60 cm. altus, plerumque circa 40 cm., strictus
vel infra spicam flexuosus. Vaginae basales 2, obtusae, glabrae, 2-5-5 et 5 -12 cm -
longae. Folia plerumque 3, vaginantia laminis acuminatis involutis 7*5-1 -5 cm.
longis, circa 1-5 mm. latis; folium infimum ad 28 cm. longum, supremum
4-5-9 cm. longum. Spica 1-3 - flora, floribus odoratis rosaceis vel pallide
rosaceis, raro subalbidis, segmentis inferioribus basin versus coccineo-maculosis.
Bractea lanceolata, acuminata, 3-5-5 cm. longa, 7-9 mm. lata; bracteolae
2-4-3-8 cm. longae, minute bicuspidatae. Perianthii tubus parum curvatus,
4-4-8 cm. longus, supra subcylindricus, basi 1-5-2 mm. diam., gradatim
ampliatus, in faucibus 0-8-1-2 cm. diam.; segmenta subaequalia vel supremum
leviter latius, obovata, obtusa vel acuta, patentia apicibus reflexis, 2-2-3 cm -
longa, 1-3-1-7 cm. lata, supremum ad 2 cm. latum. Stamina arcuata, e perianthii
tubo brevitur exserta, fllamentis 2-5-3 cm. longis antherisque 1—1-2 cm. longis.
Stylus apicem antherarum attingens vel ad 5 mm. ultra, ramis circa 4 mm.
longis, apice stigmatifero orbiculari ad 2-5 mm. lato. Ovarium 6-7 mm. longum,
2-5-3 mm - diam.
Hab. Cape Province. Galedon Division: south slopes of mountains near
Danger Point, Linley, S. A fir. Mus. No. 60214 (Type, in S.A.M. Herb.), 28 April
1948; Bredasdorp Division: slopes of hills above Pearly Beach, between
Danger Point and Quoin Point, Lewis, S. Afr. Mus. No. 60882 (S.A.M. Herb.),
June 1950; near Bredasdorp, Judd, S. Afr. Mus. No. 55622 (S.A.M. Herb, and
s. n. in Bolus Herb.), May 1940.
Corm globose, 1-1-5 cm - diam.; tunics brown, finely reticulate. Stem slender,
30-60 cm. high, usually about 40 cm., 1-5-2 mm. diam. at the base, straight or
flexuose below the spike. Basal sheaths 2, glabrous, 2-5-5 anc * 5-12 cm. long.
Leaves usually 3, the lowest up to 28 cm. long, the uppermost 4-5-9 cm. long,
all sheathing with short, involute, acuminate blades from 7-5-1-2 cm. long,
about 1*5 mm. wide. Spike 1-3 - flowered; flowers sweet-scented, pale or deep
pink, rarely whitish, the lower lobes mottled with crimson near the base. Bract
lanceolate, acuminate, 3-5-5 cm. long, 7-9 mm. wide; bracteoles 2-4-3-8 cm.
128
ANNALS OF THE SOUTH AFRICAN MUSEUM
Figure 5. Gladiolus meridionalis Lewis. 1. Flowering plant, two- thirds natural size. 2. Bract,
two-thirds natural size. 3. Bracteoles, two-thirds natural size. 4. Flower, front view — two-thirds
natural size. 5. Longitudinal section of flower — two-thirds natural size. 6. Top of style and style
branches, X 2. Del. G. J. Lewis.
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES 1 29
long, minutely bicuspidate. Perianth with tube slightly curved, 4-4-8 cm. long,
widening gradually from 1-5-2 mm. diam. at the base to 0-8-1-2 cm. diam. at
the throat, the upper half subcylindrical ; lobes subequal or the uppermost
slightly wider than the others, obovate, obtuse or acute, all spreading with
reflexed tips, 2-2-3 cm - long, 1-3-1-7 cm. wide, the uppermost up to 2 cm. wide.
Stamens arcuate, very shortly exserted from the perianth tube; filaments 2-5-3 cm -
long; anthers 1—1-2 cm. long. Style reaching to the top of the anthers, sometimes
extending 5 mm. above them; style branches up to 4 mm. long, with flat,
orbicular or slightly retuse stigmas up to 2-5 mm. wide. Ovary 6-7 mm. long,
2-5-3 mm « diam.
This species is confined to a fairly small area near the coast in the southern-
most part of the continent, between Danger Point in the Caledon Division and
Cape Agulhas in the Bredasdorp Division. Combined with the delicate colour-
ing of the flowers, it has a strong, sweet scent, like that of G. maculatus Sw., the
'Small Brown Afrikaner', to which it is very closely allied. It is distinguished
from G. maculatus by its larger flowers which are pink instead of yellow-brown,
its much broader perianth lobes and slightly longer and more cylindric perianth
tube. Both species flower during the autumn or early winter months, from the
end of April to June.
In the majority of species of Gladiolus the upper third or half of the perianth
tube widens abruptly to the throat, but there are several exceptions, such as
G. debilis, G. tristis, G. maculatus and others, including the species described and
figured here, in which the perianth tube widens gradually from the base to the
throat, the upper half being more or less cylindrical. The closely allied genus
Homoglossum is separated from Gladiolus only on account of the structure of the
perianth tube which is narrowly cylindric in the lower half and broadly cylindric
in the upper half. G. meridionalis is close to the border-line between Gladiolus
and Homoglossum and it is interesting to note that G. maculatus, to which it is so
closely related, is known to hybridize with Homoglossum Priorii when these two
occur in the same locality; both have the same flowering season and there are
several records of hybrids.
N. E. Brown, after examining the types of the South African Iridaceae
described by N. L. Burmann in 1 768 in his Florae Capensis Prodromus, published
a paper on the results of his comparisons (Kew Bull. (1929) 129-139). He made
several new combinations and cleared up some misconceptions but also
remarked that he had failed to match some of Burmann's specimens with any
at Kew. When I examined Burmann's specimens in the Herbarium at Geneva
in 1949 (by courtesy of Prof. C. Baehni, Director of the Botanic Gardens) I
found that the position regarding two of the specimens which have been the
cause of some confusion, and about which N. E. Brown had expressed some
doubt, could be clarified.
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
1. Antholyza caryophyllacea Burm. f. Brown stated that he could not match
this plant with any species at Kew. He added that it was a true 'Antholyza',
with entire (not bifid) style arms and not at all like Watsonia humilis Mill, to
which Baker had referred it as a synonym. In 1932, in a revision of the species
previously erroneously ascribed to Antholyza, he transferred this species to
Homo glos sum, as H. caryophyllaceum (Burm. f.) N. E. Br. {Trans. Roy. Soc. S.A.,
xx, 279).
This species was not known in South Africa so I examined Burmann's specimen
with considerable interest, but found that it was not possible to agree with
N. E. Brown in regarding it as a Homoglossum as the plant is one of the forms of
Gladiolus hirsutus Jacq. The leaves of Burmann's specimen are ensiform, with
prominent veins and cartilaginous margins, and are glabrous except for a few
rather inconspicuous hairs on the ventral margins in the lower sheathing part.
His plant matches Ecklon No. 156 (at Geneva) but differs from MacOwan
No. 585, from Groenkloof (Geneva, Kew, Bolus Herb, and S.A.M. Herb.), in
having a slightly longer and narrower perianth tube. Gladiolus hirsutus is a very
variable species, as was noted by Ker in 1802 : Tt scarcely ever blows two years
together of the same size or colour' {Bot. Mag., t. 574). The size of the plants
in the natural habitat is extremely variable, those growing in dry, open places
frequently being dwarfed and stunted, while others from the same area, growing
in moister or more shaded places, are tall and robust. The pubescence on the
leaves is also variable, the blades of the leaves in many specimens being glabrous,
with only a few hairs on the sheath, as in Burmann's specimen. Another
variable feature is the perianth tube, both in length and shape; in some
specimens it is almost cylindrical, as in the genus Homoglossum (cf. Burmann's
specimen), while in others it is comparatively shorter and broader, with the
upper half distinctly funnel-shaped (cf. MacOwan 585), and there are inter-
mediate forms.
The combination Gladiolus caryophyllaceus (Burm. f.) Poir. (in Lam. Encyc.
Suppl. ii, 795) was published in 181 1, and since this name is correct and valid it
should be restored to this species, with synonymy as follows: Antholyza caryo-
phyllacea Burm. f., Prodr. Fl. Cap., 1 (1768) ; Houtt., Nat. Hist., II, 12: 63, t. 79,
f. 3 (1780); Antholyza caryophyllea Panzer, Pflanzcnsyst., 1 1 : 76 (1784); Gladiolus
hirsutus Jacq., Coll., iv, 161 (1790); Ic, t. 250 (1786-93); Red., Lil., t. 273
(1809); Ker, Gen. Irid., 132 (1827); Baker, Handbk. bid. (1892) and FL Cap.,
vi, 153 (1896); G. hirsutus Jacq. var. roseus Ker, Bot. Mag., t. 574 (1802);
G. roseus Andr., Bot. Rep., t. 11 (1797); G. similis Eckl., Top. Verz*, 40 (1827);
Homoglossum caryophyllaceum (Burm. f.) N. E. Br., Trans. Roy. Soc. S.A., xx, 279
(i932).
The paragraph by Poiret stating that this species was near to Antholyza
Cunonia {—Anomalesia cunonia (L.) N. E. Br.) is misleading, as Burmann's plant
does not in any way resemble Anomalesia cunonia.
2. Gladiolus pyramidalis Burm. f. Concerning this species N. E. Brown noted:
'No specimen so named. But there is a specimen from which a name on the
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES 131
bottom of the sheet has evidently been cut off, and which bears on one label
with it the name i( Watsonia pyramidata et spicata" and on another label "Ixia
foliis nervosis fl. pyramidali tubuloso cameo". It is a species of Watsonia allied to
W. brevifolia Ker, but I cannot match it at Kew. It agrees very well with the
description of G. pyramidalis Burm. As I find that some of the names under which
Burmann described his plants have been crossed out, and other (sometimes
unpublished) names substituted, it is quite possible that if this is the type of
G. pyramidalis Burm., the name for some reason may have been removed from
the sheet.'
Among Burmann' s specimens I found one which is named by him G. pyra-
midalis; N. E. Brown, although he saw this specimen and named it 'Tritonia',
must have overlooked Burmann's name, as is evident from his note. The plant
is a dwarf specimen (about 6 to 7 in. high) of Ixia patens Ait., but with large and
normal flowers, the bracts nearly half an inch long, the perianth tube not
exceeding the bracts and the lobes about one inch long. It agrees with Burmann's
description which is very brief, but the last four words 'tubo lineari, longitudine
spathae' leave no doubt that the specimen named G. pyramidalis in his collection
is the one he described, and is the type of his species. The description of the
perianth tube cannot possibly apply to any species of Watsonia.
As Lamarck described a species of Lapeyrousia under the name Ixia pyramidalis
[Encyc, hi, 334 (1789) ), this name is not available for Burmann's plant and
Gladiolus pyramidalis Burm. f. should be added to the list of synonyms of Ixia
patens Ait.
The confusion regarding the identity of Gladiolus pyramidalis probably arose
less than twenty years after the name was first published, when Lamarck
described a species of Watsonia under this name (Encyc, ii, 726 (1786) ), and
remarked ( an Glad, pyramidalis Burm. Prodr. 2 sedflores non imbricati\ I have seen
Lamarck's specimen, named by him, and it is the species now known as
Watsonia pyramidata (Andr.) Stapf (W. rosea Ker). It is fairly evident that
Lamarck, like N. E. Brown, must have considered that the species of Watsonia
referred to by Brown was the type of Burmann's Gladiolus pyramidalis.
Gladiolus pubescens Lam., Illus. I, 119 (1 791)
The use of this name by Lamarck has not been recorded in the Kew Index,
although Lamarck published a description of the species and there is a specimen
bearing his name in his herbarium in Paris. On examining Lamarck's type
(through the courtesy of Prof. Humbert, Director of the Department of Botany
of the Museum National d'Histoire Naturelle) I found that it is a very distinct
species of Babiana, with plicate, truncate, cuneate leaves. It is the same as
Drege 2627, the type of Babiana cuneifolia Baker, but as Lamarck's specific
epithet has priority I propose to reinstate it, so that this species now becomes
Babiana pubescens (Lam.) Lewis, comb. nov. (Syn. Gladiolus pubescens Lam.,
Illus., I ( 1 791) 119; Babiana cuneifolia Baker, Jour n. Bot., 1876, 335; FL Cap., vi
(1896) no).
132 ANNALS OF THE SOUTH AFRICAN MUSEUM
Type, without locality, De Vaillant in Herb. Lamarck (Paris).
This species has been collected in three or four localities in Namaqualand
and was figured by myself in 1938 in Flowering Plants ofS. Africa (xviii, p. 685).
There are at present two species of Gladiolus named G. pubescens, i.e. G.
pubescens Baker and G. pubescens Pax, but as the name is not valid for either of
them they will both have to be renamed, and I propose to name them as follows :
Gladiolus pubigerus Lewis nom. nov. (G. pubescens Baker, Journ. Bot., xiv
(1876) 333; FLCap.,vi, 142 (1896).).
Gladiolus pubescifolius Lewis nom. nov. (G. pubescens Pax, Engl. Bot.
Jahrb., xv (1893) 154; Baker in Fl. Trop. Afr., vii (1898) 364).
Moraea unguicularis Lam., 77/. Gen. No. 490 (1791); Encyc, iv, 274 (1797).
The specimen bearing this name in Lamarck's herbarium, collected by
Sonnerat, is definitely not a Moraea, and it is difficult to understand how it came
to be placed in this genus. It is an Exohebea and is the species which N. E. Brown
described as Gladiolus fraternus in 1928 (= Exohebea fraterna (N. E. Br.) Foster).
On Lamarck's sheet there is the following note: 'An Ixia aut Moraea. Germ,
inf. stam. 3. stigma 3 fidum. e Cap. B. Spei. rad. est bulbus subrotundus'.
Lamarck gave a fairly detailed description of the plant in his Encyc, iv, p. 275,
and there is no doubt that the specimen so named in his herbarium was the
one described. As Lamarck's specific name has priority I propose to retain it
for this species of Exohebea, and name it
Exohebea unguicularis (Lam.) Lewis, comb. nov. (Syn. Moraea unguicularis
Lam., III. Gen. No. 490 (1 791) 114; Encyc, iv/274 (1797); Vieusseuxia ungui-
cularis Roem. & Schult., I, 491 (181 7), ex parte (excl. syn. and description of
M. unguiculata Ker) ; Gladiolus fraternus N. E. Br., Journ. Linn. Soc, xlviii, 26
(1928) ; Exohebeafraterna (N. E. Br.) Foster, Contrib. Gray Herb., cxxvii, 37 (1939)).
Type, without precise locality, Sonnerat, in Herb. Lamarck (Paris).
Roemer and Schultes evidently thought that Moraea unguiculata Ker was the
same as M. unguicularis Lam., but this is incorrect as there is no connection
between these species. M. unguiculata Ker is a Moraea and it was figured and
described as a new species in Bot. Mag., t. 593 (1802), as is clear from the fact
that Ker made no reference to M. unguicularis Lam., and from his statement:
'Introduced from the Cape by Mr. Alderman Hibbert, at whose garden our
drawing was taken, and where alone, we believe, it is at present to be found.'
Gynandriris torta (L. Bol.) Foster.
This combination, made by Foster in Contrib. Gray Herb., cxiv, 41 (1936), is
correct and should remain as it is. In making this new combination he followed
T. T. Barnard, who transferred Moraea torta L. Bol. to the genus Helixyra {Iris
Trbk. (1932), 52), though, as was pointed out later by Foster, the generic name
established by N. E. Brown was invalid and the proper name for the genus is
Gynandriris. In a later publication {Contrib. Gray Herb., cxxvii (1939) 48), Foster
referred this species back to the genus Moraea, but this is incorrect. In doing so
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES 1 33
he stated that since making the combination three years earlier he had studied
the description of Moraea torta and the figure which accompanied it; unfor-
tunately, however, both are misleading, as there is no mention of the prolonga-
tion of the ovary in the description, nor is it shown in the figure. In this species
there is a sterile prolongation of the ovary from 2 to 2-5 cm. long.
It is very probable that Moraea Pritzeliana Diels {Engl. Jahrb., xliv (1909) 117),
from the Galvinia Division, is the same species, as far as can be judged from the
description, but as there is no mention at all of the ovary in Diel's description
it is not possible to come to any decision without examining his type (if it is still
in existence) . If it proves to be conspecific with Gynandriris torta then Diel's
specific name has priority.
Lapeyrousia divaricata Baker
In the Journ. S. Afr. Bot., vii, 56 (1941), I placed Chasmatocallis Macowani
Foster under Lapeyrousia setifolia (L. f.) N. E. Br. In doing so I had accepted
N. E. Brown's name for the species and had overlooked Foster's note in Contrib.
Gray Herb., cxiv, 48 (1936), in which he pointed out that Brown's combination
was not valid. The correct name for this species is Lapeyrousia divaricata Baker
{Journ. Bot., xiv, 337 (1876); Fl. Cap., vi, 91 (1896). Syn. Gladiolus setifolius
L. f., Suppl., 96 (1781); Thunb., Diss. Glad., 18 (1784); Lapeyrousia setifolia
(L. f.) N. E. Br., Journ. Linn. Soc, xlviii, 30 (1928); Chasmatocallis Macowani
Foster, Contrib. Gray Herb., cxxvii (1939) 40).
Geissorrhiza teretifolia Lewis
It is regretted that there has been some confusion about this species due to
my own error in citing a specimen of G. geminata when I described the species
{Journ. S.A. Bot., vii (1941), p. 48). Since the publication of my description and
Foster's Revision of Geissorrhiza {Contrib. Gray Herb., cxxxv (1941) 1-78) I have
examined the type of G. geminata and found that Foster was quite right in
regarding this specimen, Lewis in Herb. Bolus No. 20414, as being identical
with G. geminata, and the reference to it should be deleted from my original
description. The other specimens cited, which unfortunately were not seen by
Foster, all belong to G. teretifolia, a species quite distinct from G. geminata E. Mey.
ex Baker.
At the end of September 1951 the locality near Brand Vlei in the Worcester
Division, where the specimen Lewis in Herb. Bolus No. 20414 was collected,
was revisited, and more material of G. geminata collected. Owing to my error
Foster applied my description of the corm of G. teretifolia to the corm of G. gemi-
nata, as there was no complete corm on the type, but the correct description of
the corm of G. geminata can now be given, as follows : Corm ovoid, up to 9 mm.
diam., usually 6-7 mm., with brown concentric tunics which are very shortly
cusped at the apex.
The corm tunics of G. teretifolia are entirely different, being very dark, almost
black in colour, imbricate and split upwards from the base. The basal part of
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
the stem in this species is a dark red-purple colour and the single branch, when
present, arises from below the middle of the stem, the cauline leaf which sub-
tends it having a fairly long, terete blade, equal to or up to twice as long as the
slightly ventricose striate sheath. The flowers are more numerous and larger
than those of G. geminata. In G. geminata the stem is white to greenish at the base,
the cauline leaves are comparatively shorter, with the sheath not ventricose or
striate, and the branches (one to three) arise above the middle of the stem. Both
species grow in damp, marshy places, in shallow water on the edge of seasonal
marshes and pools.
G. teretifolia belongs to Foster's subsection Ventricosae; it is very closely allied
to G. imbricata (De la R.) Ker var. Brehmii (Klatt) Foster, and on comparison
might even prove to be the same, though the leaves of the variety are described
by Foster as one-nerved, whereas the leaves of G. teretifolia, as seen in the living
plants, are distinctly terete, 4-grooved, and slightly fleshy.
Geissorrhiza furva Ker ex Baker
There is some uncertainty about the identity of this species and it is doubtful
whether Foster's treatment of it in his Revision of Geissorrhiza is correct. Baker,
when he first described it (Handbk. hid. (1892) 155), cited three specimens,
Masson, Drege 8478, and Bolus 4341 ; in the Flora Capensis (vi, 70) he added
another, Ecklon and £eyher 217. Foster, in his revision of the genus, designated
Drege 8478 as the type (Contrib. Gray Herb., cxxxv (1941) 26), and remarked:
'Of the three specimens cited in the original description, I have seen only one.
Since this fits the description reasonably well, and even has the remnant of a
corm, it may be designated as the type. Baker stated that the anthers were longer
than the filaments but in the type they are only half as long.'
It is evident that Foster did not see Masson's specimen, since he ha> not
mentioned it in his revision, but he must have seen Bolus 4341 as he cited this,
among specimens seen by him, under G. monantha (p. 40) . {Ecklon and £eyker 217,
according to Foster (p. 22) is G. juncea.) Ker's name, although a nomen nudum,
was published in 1804 (Koenig and Sims, Ann. Bot., i, 224), so there can be no
doubt that Masson's specimen was the only one of those cited by Baker which
was seen by Ker, and must be the one which Ker intended to bear the name.
Whether or not it is the one on which Baker's description was based cannot
be decided without seeing Masson's plant. From Baker's description it seems
very probable that the plant he described is the same as G. monantha (Thunb.)
Eckl., but his description might perhaps have been drawn from Bolus 4341,
which is that species.
The flowers o^G. furva were described by Baker, and also by Foster, as bright
red-purple. On examining Drege 8478, collected between Paarl and Pont,* I
found that it does not agree even 'reasonably well' with Baker's descriprion, for
in addition to the difference in the stamens noted by Foster, the colour of the
flowers is quite different, those of Drege 8478 being a bright golden-yellow,
* Melck's Pont, Berg River.
IRIDACEAE NEW SPECIES AND MISCELLANEOUS NOTES 1 35
nearly always changing to a dingy brownish colour when dried, as in Drege's
specimen. The following specimens in the Bolus Herbarium are the same as
Drege 8478 : Paarl Division : flats north of Paarl, Leighton 2002 ; same locality,
Esterhuysen 6127; north base of Joostenberg, Pillans 9262; Wellington, Dawson
(B.H. No. 14012); flats north of Hercules Pillar, between Mulders Vlei and
Joostenberg, Leighton 555; Hercules Pillar, Compton 160 14.
In my opinion the question regarding the correct identity of Geissorrhizafurva
can only be decided by comparing Drege 8478 with Masson's plant. If these
prove to be conspecific then it will be necessary to change Foster's description
of the colour of the flowers from 'red-purple' to bright yellow. If, however, they
are not the same, then Drege 8478 (and the specimens in the Bolus Herbarium
which match it) requires a new name. If Masson's plant proves to be the same
as Bolus 4341, i.e. G. monantha, then G.furva Ker ex Baker should be treated as
a synonym of G. monantha (Thunb.) Eckl.
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XL
PART IV, containing: —
5. A revision of the Genus Synnotia. By G. J. Lewis, B.A., Ph.D.
ISSUED APRIL 1956 PRICE 3s.
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TRUSTEES OF THE SOUTH AFRICAN MUSEUM
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5- A revision of the Genus Synnotia. By G. J. Lewis, B.A., Ph.D.
Introduction and Historical Note
The history of the first species found at the Cape goes back more than two
centuries, to 1 739, when Breynius described and figured the species which was
named Gladiolus villosus by Burmann f. in 1 768. In 1 784 this same species was
described and figured by Thunberg as Gladiolus bicolor. Sweet established the
genus Synnotia in 1826, with an illustration and description of Synnotia variegata,
and the following year he transferred Gladiolus bicolor to Synnotia, retaining
Thunberg's specific name; from that date until 1929, when the older specific
name was restored by Dr. N. E. Brown, the species was known as Synnotia
bicolor. It was evidently fairly well known in Europe by the end of the eighteenth
century, and in the first half of the nineteenth century S. villosa, and, after its
discovery, S. variegata as well, became popular garden plants in England and
Europe, judging from the illustrations published in botanical and horticultural
works of that period. In England figures of one or both species appeared in the
Botanical Magazine, Sweet's British Flower Garden and Ornamental Flower Garden
and Loudon's Ladies' Flower Garden, while in Europe Thunberg and Jacquin
published illustrations of S. villosa before the end of the eighteenth century.
Another species, illustrated by Jacquin and named by him Gladiolus galeatus,
was named Synnotia galeata by Sweet, but after that little seems to have been
known about it and it does not appear to have been rediscovered in the field
until about a century after the publication of Jacquin's figure. Although
Durand and Schinz retained it as a distinct species in their Conspectus Florae
Africae, vol. v (1895), Baker evidently considered it to be a form of S. villosa,
for he placed it as a synonym of S. bicolor in the Flora Capensis, and in this
country it has remained confused with that species until now. In this revision
it is restored to its proper status as a distinct species.
In spite of its early history very little herbarium material of the genus seems
to have been collected before 1896, when Baker's work on the Iridaceae was
published in vol. vi of the Flora Capensis. Baker himself saw and cited only
seven specimens, which he placed under two species and one variety. In 1901
he described a third species but this has to be excluded from the genus as it is
a mixture of Synnotia and Sparaxis (see note on S. stenophylla Bak. at the end of
this paper). Except for the publication of a new species by Dr. L. Bolus in 1923,
no work has been done on the genus during the past sixty years, although a
great deal more material has been collected, and it is not surprising to find that
Baker's work, based on his meagre knowledge of the genus and the few speci-
mens available to him, is inadequate and requires revision. This indeed is
true of most of the genera of Iridaceae described in the Flora Capensis.
137
VOL. XL. PART 4 JUN 4 l95fc
I38 ANNALS OF THE SOUTH AFRICAN MUSEUM
The present revision is based on a study of the plants in the field and an
examination of eighty-two collectings, the majority of which are housed in
herbaria in South Africa. Unfortunately it has not been possible to examine
all the material in herbaria in England and Europe, though some of it has
been seen.
For the loan of specimens and for facilities granted me to examine material
in their herbaria sincere thanks are expressed to the Directors and Curators
of the following institutions:
Bolus Herbarium, University of Cape Town ; National Herbarium, Pretoria ;
National Botanic Gardens, Kirstenbosch ; Botanical Museum, Uppsala;
Conservatoire de Botanique, Geneva; Botanical Museum, Zurich; Musee
d'Histoire Naturelle, Paris.
Geographical Note
Synnotia is a small genus, endemic in South Africa where it is confined to a
few districts in the south-western part of the country, from the northern area
of the Cape Peninsula northwards to the Van Rhynsdorp Division, with one
record from the Calvinia Division. With the exception of the record from near
Calvinia, all the others are from low or fairly low altitudes, on flats, hills or
lower mountain slopes in the coastal districts, the Olifants River Valley and the
base of the escarpment below Van Rhyn's Pass. (See map.)
S. Metelerkampiae L. Bolus (treated as a variety of S. variegata in this revision)
has been recorded from the western slopes of the Cedarberg, up to an altitude
of 2,000 ft., but the plants found at these higher altitudes are obviously dwarfed
and stunted compared with those from lower altitudes. The most easterly
record and the highest is one of S. galeata, from about 1 2 miles south-west of
Calvinia, at an altitude of about 3,000 ft.
The genus Sparaxis is very closely related to Synnotia, from which it differs
mainly in having more regular flowers, with equal or subequal perianth lobes.
This also is a small and endemic genus, with much the same range of distri-
bution as Synnotia, though it does not go further north than the Clanwilliam
district and extends further to the south-east, to the Caledon and Swellendam
Divisions. Like Synnotia it occurs at low altitudes, where in some parts it is
extremely common, but is not found on the higher mountain slopes.
Description of the Genus
Synnotia
Sweet, Brit. Flow. Gard., t. 150 (1826); Baker, Fl. Cap., vi, 134 (1896);
Anactorion Raf., Fl. Tellur., iv, 34 (1836).
Named in honour of W. Synnot who collected a number of plants at the
Cape.
Corms small to medium sized, globose or ovoid, with pale brownish tunics
of short, hard reticulate fibres or long fine hair-like fibres extending up in a
A REVISION OF THE GENUS SYNNOTIA
139
neck. Basal sheaths 2, acute or acuminate, membranous and colourless or the
upper sometimes green-tipped. Leaves 5 to 9 in a fan-like distichous rosette,
ensiform or narrow-ensiform, acute, subacute or obtuse, glabrous, closely
veined, the veins seldom conspicuous; cauline leaves 1 to 3, acute or acuminate.
Stem erect, glabrous, simple or branched. Inflorescence spicate, the spikes laxly
2 to 7-flowered, or occasionally the flowers solitary. Bracts membranous, ovate,
cuspidate and irregularly denticulate, with numerous reddish-brown vein-like
streaks above the base; bracteoles similar and almost as long as the bracts,
united nearly to the top, bicuspidate with acute to setaceous-acuminate cusps.
Flowers distinctly zygomorphic; perianth tube short or long, the lower half or
more cylindrical, curved above and becoming gradually or abruptly infundi-
buliform; perianth lobes obtuse or rarely subacute, the uppermost larger than
the others, erect in fully open flowers, the 3 lower connivent, forming a more
or less projecting reflexed lip. Stamens contiguous, arcuate, inserted at the
base of the expanded part of the perianth tube, usually reaching to the middle
of the uppermost perianth lobe; filaments filiform; anthers linear or linear-
oblong, attached shortly above the base. Ovary small, oblong or ovate-oblong;
style reaching to the middle of the anthers, with short entire stigmas either
expanding from near the middle upwards and conduplicate or slightly
expanded at the apex only. Capsules subglobose or turbinate. Seeds fairly
numerous, comparatively large, smooth and shining, rounded or slightly
angled by pressure.
The type species is S. variegata Sweet, Brit. FL Gard., t. 150.
Key to the species
1. Flowers small, not more than 2-5 cm. from base of ovary to top of
uppermost perianth lobe. . . . 1. parviflora
1. Flowers not less than 3 cm. from base of ovary to top of uppermost
perianth lobe, usually considerably more.
2. Cylindrical part of perianth tube up to 1 cm. long, rarely slightly
longer.
3. Corm tunics of hard reticulate fibres not extending up in a neck;
lower side perianth lobes auriculate above the base; uppermost
lobe more or less obovate. ... 2. villosa
3. Corm tunics of numerous fine matted fibres extending up in a
neck; lower side perianth lobes not auriculate; uppermost lobe
not obovate, up to twice as long as wide.
4. Cylindrical part of perianth tube not more than 7 mm. long;
uppermost perianth lobe of mature flowers usually slightly
more than twice as long as wide. . . . 3. galeata
4. Cylindrical part of perianth tube not less than 1 cm. long,
usually more; uppermost perianth lobe up to twice as long as
wide. ... 4. Roxburghii
2. Cylindrical part of perianth tube 2 cm. or more long.
5. Corm tunics of fine hair-like fibres extending up in a neck;
cylindrical part of perianth tube seldom more than 2 cm. long,
curved above but not geniculate. ... 4. Roxburghii
140 ANNALS OF THE SOUTH AFRICAN MUSEUM
5. Corm tunics of hard reticulate fibres not extending up in a neck;
cylindrical part of perianth tube 2-5 cm. long or more, geniculate
at the top.
6. Upper part of perianth tube broadly funnel-shaped, 1 to 1-5
cm. diam. at the top; length from bend in perianth tube to
top of uppermost lobe 3 to 3*5 cm. . . . 5. variegata
6. Perianth tube usually 0-5 to o-8 cm. diam. at the top, rarely
up to 1 cm.; length from bend in perianth tube to top of
uppermost lobe rarely more than 2-5 cm. . . . 5. variegata var.
Metelerkampiae
i . S. parviflora Lewis sp. nov.
Cormus ovoideus, 1-2-1-5 cm. diam., tunicis e fibris numerosis tenuibus
compositis. Caulis plerumque 15-30 cm. altus, simplex vel basin versus uno
ramo. Folia 7-9, acuta, 3-18 cm. longa, 0-3-1-2 cm. lata, nervis inconspicuis.
Spica 1-4-flora, plerumque 2-3. Bractea 1-1*5 cm - longa, o-8-i cm. lata, plus
minusve tricuspidata, lineis rubrido-brunneis distincta; bracteolae bractea
aequilongae, cuspidibus acutis vel acuminatis. Flores gilvi extrinsecus lilacino-
suffusi, segmentis inferioribus lateralibus croceis basi minute brunneo-
maculatis. Perianthii tubus 9 mm. longus, parum curvatus, parte cylindrica
6 mm. longa, parte superiore leviter infundibuliformi ; segmenta obtusa, 1-1 -2 cm.
longa, 5-7 mm. lata, supremum quam cetera leviter longius et latius, 3 inferiora
basin versus breviter attenuata. Filamenta 6 mm. longa; antherae 4 mm. longae.
Ovarium 4 mm. longum, 2-5-3 mm. diam.; styli rami 2 mm. longi.
Corm ovoid, 1-2-1-5 cm. diam.; tunics of very fine matted fibres. Stem
usually 15-30 cm. high, simple or with 1 branch arising fairly near the base.
Leaves 7-9, acute, 3-18 cm. long, 0-3-1-2 cm. wide, the veins inconspicuous.
Spikes 1-4-flowered, usually 2-3. Bract 1-1-5 cm - l° n g- o-8-i cm. wide, more
or less tricuspidate, streaked with dark red-brown; bracteoles as long as the
bract, with acute or acuminate cusps. Flowers cream-coloured, flushed with
mauve outside, the lower side lobes bright yellow with a pair of small brownish
comma-shaped marks at the base of each; perianth tube 9 mm. long, the cylin-
drical part 6 mm. long, slightly curved and expanded above; perianth lobes
obtuse, 1—1-2 cm. long, 5-7 mm. wide, the uppermost only slightly longer and
wider than the others, the 3 lower very shortly attenuate at the base. Filaments
6 mm. long; anthers 4 mm. long. Ovary 4 mm. long, 2-5-3 mm. diam.; style
branches 2 mm. long, the upper half flattened, conduplicate and recurved.
Type. Lewis (65637 in the S. Afr. Mus. Herb.).
Malmesbury Division: between Mamre and Darling, on sandy ground,
Lewis (S.A.M. 65637) ; near Darling, Lewis (S.A.M. 56745) ; near Hopefield,
Schlechter 5304; Bolus (B.H. 12853); Langebaan, Salter 3025; Lewis (N.B.G.
2032/32 in Bol. Herb.).
Flowering season. September.
A very distinct species. The flowers are by far the smallest in the genus and
the perianth lobes less irregular than in the other species. Its nearest ally is
S. villosa from which it differs in having finer corm tunics and smaller flowers,
A REVISION OF THE GENUS SYNNOTIA
141
Figure 1. Synnotia parviflora Lewis. 1. Plant in flower. 2. Bract. 3. Rracteoles. 4. Flower, laid
open. 5. Stamen, x 2. 6. Gynaeceum, x 2. Del. G. J. Lewis.
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
with the uppermost lobe only slightly larger than the others and the lower
side lobes not auriculate.
Among the specimens in the Bolus Herbarium collected by Bolus (B.H.
no. 12853) there is one very robust plant, about 50 cm. high, with leaves about
i*5 cm. wide, but with the normal small flowers. All the others on the same
sheet are the typical rather small plants described above.
2. S. villosa (Burm. f.) N.E. Br., Kew Bull., 1929, 133; Gladiolus villosus
Burm. f., Fl. Cap. Prodr., 2 (1768); Gladiolus bicolor Thunb., Diss. Glad., 16,
t. 2, f. 1 (1784) ; Prodr. 8 (1800) ; Jacq., Ic, ii, t. 240 (1786-93) ; Coll. SuppL, 25
(1796) ; Ixia bicolor Sims, Bot. Mag., t. 548 (1802) ; Sparaxis bicolor Ker, Konig &
Sims Ann., i, 225 (1805); Synnotia bicolor (Thunb.) Sweet, Hort. Brit., ed. 1,
398 (1827); e d- 2, 501 (1830); Klatt, Linnaea, xxxii, 750 (1863); Baker, FL
Cap., vi, 134 (1896).
Corm globose, 1-1-5 cm - diam., with hard tunics, reticulate above and the
lower half of rather coarse vertical parallel strands. Stem 12-35 cm - high?
usually slightly flexuose, simple or with 1 or 2 branches. Leaves usually 7 in
the rosette, acute or subacute, rarely obtuse, 3-21 cm. long, 0-4-1-5 cm. wide,
usually 10-18 cm. long and 1 cm. wide, the veins inconspicuous. Spike 1-5-
flowered, usually 2-3-flowered. Bract 1-5-2-5 cm. long, 1-1-4 cm - wide,
setaceous-cuspidate, the margins irregularly denticulate, sometimes more or
less cuspidate or lacerate, colourless or pale straw-coloured at the base with
reddish-brown streaks above; bracteoles similar, setaceous-bicuspidate. Flowers
pale yellow, the uppermost lobe pale mauve, sometimes the tips of the lower
lobes as well; perianth tube 1-5-2 cm. long, the cylindrical part 0-7-1 cm. long,
the upper part broadly funnel-shaped: perianth lobes usually with the lower
half more or less connivent, the uppermost erect and the other 5 with the
upper half somewhat patent or recurved; uppermost lobe obovate, 1-4-1-9 cm.
long, 0-9-1-3 cm. wide, sometimes slightly emarginate; upper side lobes
1 -1 -4 cm. long, 5-6 mm. wide; lower side lobes shortly unguiculate, auriculate
above the claw, 1—1-3 cm - l° n g> 5~8 nun. wide above the claw; lowest lobe
smaller, 7-9 mm. long, 4-5 mm. wide, not unguiculate nor auriculate.
Stamens reaching to or slightly above the middle of the uppermost lobe, slightly
arcuate; filaments about 1-5 cm. long; anthers 4 mm. long. Ovary 4-5 mm.
long, 2-5-3 mm « diam.; style branches 3 mm. long, the upper half expanded,
conduplicate, minutely ciJiate.
Lectotype. As there is no specimen in Burmann's collection and the figure
on which his name is based (Breyne, Ic. Rar. PL, t. viii, f. 2) dates back to 1739,
it seems advisable to retain Thunberg's type of Gladiolus bicolor as the type of
this species. There are two sheets in his herbarium in Uppsala, from Groene-
kloof, in the Malmesbury Division.
Cape Division: Lion's Head, MacOwan 798; Signal Hill, Marloth 188; fields at
Observatory, W. Dod 1443; between Salt River and KaJabaskraal, 13 miles
from Cape Town, Hutchinson 1 75 ; Vissershok, Barker 1 8 1 1 . Malmesbury
A REVISION OF THE GENUS SYNNOTIA 1 43
Division : Groenekloof, Thunberg ; north-east slopes of Dassenberg, Pillans 6859 ;
Malmesbury, Barker 4667, 6394; Guthrie 2386; Lewis (S.A.M. 60674, 65642);
between Mamre and Darling, Lewis (S.A.M. 56747); Darling, Compton 19880.
Piketberg Division: De Hoek, Steyn 532. Paarl Division: Paarlberg, Drege 8347;
Dal Josafat, near Wellington, Grant 2333; near Wellington, Martin, s.n.
Tulbagh Division: near Saron, Schlechter 4869; Tulbaghskloof, Ecklon and
Zeyher 121 (77*9); near Artois, Bolus 7586. Stellenbosch Division: Klapmuts,
Prior.
Without locality: Jussieu 3608; Lamarck (named Ixia aristata).
Flowering season. August to September.
It is rather unfortunate that the old and seemingly inappropriate specific
epithet used by Burmann has to be retained for this species which, like the
others, is entirely glabrous. There is no specimen in Burmann's collection and
he gave no description of the plant, merely citing Breynius' figure. According
to N. E. Brown this is the plant known as Synnotia bicolor (Thunb.) Sweet, but
as Burmann's name is older it should be called S. villosa. Brown added that
the figure of Breynius is a fairly good one, and quite unmistakable. More than
a century earlier Sims was of the same opinion, for in Bot. Mag., t. 548 (1802),
under Ixia bicolor ( = Gladiolus bicolor Thunb.) he gave as a synonym Gladiolus
villosus Burm. and cited the figure of Breynius.
Although there is no doubt that the figure is of this species, and the des-
cription given by Breynius applies to it, his use of the adjective villosus in the
description, which was adopted by Burmann for the specific epithet, is at
first rather puzzling. It is evident from the description and illustration, how-
ever, that it applies to the hair-like cusps at the apex and on the margins of
the bracts and bracteoles, and bearing in mind that at the time when he
described the species it was probably the only plant of its kind in Europe, this
character of the bracts was a striking one and distinguished this species from
all other species of Gladiolus, to which genus it was then considered to belong.
3. S. galeata Sweet, Hort. Brit., ed. 1, 398 (1827); ec *- 2 > 5 01 (^3°) 5 Dur.
and Schinz, Consp. Fl. Afr., v, 211 (1895); Gladiolus galeatus Jacq., Ic, ii, t. 258
(1786-93) and Coll. iv, 167 (1790), non Burm. f. (1768); Sparaxis galeata Ker,
Konig. & Sims Ann., i, 225 (1805) ; Synnotia bicolor Pole Evans in Fl. PI. ofS. Afr.,
v, t. 162 (1925), non S. bicolor (Thunb.) Sweet.
Corm oblong-ovate or ovoid, 1-2-2-2 cm. diam.; tunics of numerous fine
hair-like fibres, nearly always extending up in a neck 2*5-3-5 cm - long- Stem
12-35 cm - high, somewhat flexuose, simple or with a branch arising near the
base, rarely with 2 branches. Leaves 5-9, acute or obtuse, usually obtuse and
apiculate, 2-16 cm. long, 0-5-1-5 cm. wide, sometimes spotted or banded
with purple at the base, the marginal veins usually slightly thickened, the
other veins not very conspicuous. Spike 2-5-nowered. Bract 1-2-2 cm. long,
0-7-1 cm. wide, setaceous-cuspidate, the margins irregularly denticulate,
colourless or the upper half pale reddish-brown, speckled all over with fine
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
red-brown streaks; bracteoles similar, setaceous-bicuspidate. Flowers cream-
coloured more or less suffused with mauve, with purplish patches at the base
of the 2 upper side lobes, the 3 lower lobes bright yellow, sometimes tipped with
pale mauve; perianth tube up to 1*3 cm. long, the cylindrical part 6-7 mm. long,
narrowly infundibuliform above; perianth lobes very unequal, the uppermost
longer than the others, erect and well separated from the other 5 lobes when
the flowers are fully open, 1-8-2-6 cm. long, 0-6-1-2 cm. wide, shortly unguicu-
late and more or less oblong above the claw; upper side lobes patent, 1-5-2 cm.
long, 6-8 mm. wide; 3 lower lobes alike, forming a distinct deflexed lip,
1-3-1-6 cm. long, 4-5 mm. wide, usually very shortly unguiculate. Stamens
very arcuate ; filaments 2 cm. long; anthers 5-6 mm. long. Ovary 2-5-3 mm -
long, 2 mm. diam. ; style branches 4-5 mm. long, filiform, slightly expanded at
the apex.
Type. Jacquin, Ic, ii, t. 258.
Clanwilliam Division: west side of Olifants River, about 2 miles east of
Claypan and 7 miles south-west of Klaver, stony koppie on farm Driefontein,
Nieuwoudt (S.A.M. 67740); Claypan, Ponder (N.B.G. 1466/28 in Bol. Herb.).
Van Rhynsdorp Division: near Van Rhynsdorp, Rood, N.H. 2860 (Pretoria);
N.B.G. 2306/23 (Bol. Herb.) ; between Van Rhynsdorp and Clanwilliam,
Marloth 7469; foot of Van Rhyn's Pass, in hard reddish soil, Lewis (S.A.M.
60136); Barker 6445. Calvinia Division: Driefontein, south-west of Calvinia,
Marloth 10426.
Flowering season. End of July to middle of September.
No specimens were seen by Baker, who placed Synnotia galeata Sweet and
Gladiolus galeatus Jacq. as synonyms under Synnotia bicolor in the Flora Capensis
(vi, 135), in spite of Jacquin's excellent figure which depicts a plant quite
unlike the one figured by him as Gladiolus bicolor in the same volume {Ic, t. 240).
The two species are quite distinct, with very different corm tunics and flowers.
The corm tunics of S. villosa are of hard, reticulate fibres, not extending up
in a neck, while those of S. galeata are fine, soft and thickly matted, and nearly
always extend up in a neck to ground level. In S. villosa all the perianth lobes
are more or less connivent in the lower half, giving the flower a somewhat
closed and broadly funnel-shaped appearance, whereas in S. galeata, though
the immature flower somewhat resembles that of S. villosa, the mature flower
has a wide open appearance, with the uppermost lobe well apart from the
others and the upper side lobes also widely separated from the 3 lower. In
addition the lower side lobes of S. villosa are auriculate near the base and the
uppermost lobe is obovate, whereas in S. galeata the lower lobes are not
auriculate and the uppermost, which elongates as the flower matures, is
finally much longer and comparatively narrower, at least twice as long as
wide. For comparison a reproduction of the inflorescence of S. villosa (from
Bot. Mag., t. 548) and S. galeata (from Jacq., Ic, t. 258) is given in figure 2,
nos. 1 and 2.
A REVISION OF THE GENUS SYNNOTIA
145
Figure 2. 1. Synnotia oillosa (Burm. f.) N.E. Br., from Bot. Mag., t. 548 (Ixia bicolor Sims).
2. Synnotia galeata Sweet, fromjacq., Ic, ii, t. 258 (Gladiolus galeatus J acq.).
The purple mottling near the base of the leaves does not appear to be a
constant feature in S. galeata. The plant figured by Jacquin shows no trace
of any marks and on only two of eight specimens seen from near Claypan on
the west side of the Olifants River are there rather faint purple spots. In the
majority of specimens from other localities, however, the leaves are con-
spicuously spotted or banded with purple, as is shown on the plant figured in
Flowering Plants of S. Africa (v, t. 162). The leaves are often obtuse, usually
obtuse and apiculate, more or less as shown by Jacquin, and are not all very
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
acute as described in Flowering Plants. As well as having most of the leaves
marked with purple, plants from the foot of Van Rhyn's Pass and near Calvinia
have slightly larger and firmer bracts than those from near Claypan, but they
are not considered to be more than a local form. It is very probable that
Jacquin's type came from the west side of the Olifants River as the plants
recently collected on the farm Driefontein, near Glaypan (Sept. 1955), most
closely match his figure, and this locality was on the old route to Namaqualand
followed by the early travellers, near the Company's Drift where the Olifants
River was crossed.
4. S. Roxburgh!! (Baker) Lewis sp. nov.
5. bicolor (Thunb.) Sweet var. Roxburghii Baker, Handbk. Irid., 198 (1892);
FL Cap., vi., 135 (1896).
Cormus globosus, 2-5-3 cm. diam. tunicis e fibris copiosis capillaribus com-
positis in collum 7-8 cm. longum productis. Caulis 28-42 cm. altus, simplex
vel uno ramo interdum 2 ramis. Folia 7-9 basalia, 5-23 cm. longa, 0-5-1-4 cm.
lata, acuta vel subacuta, nervis conspicuis sed non crassis; folium caulinum 1,
4-6 cm. longum, acutum vel acuminatum. Spica 2-7-flora. Bractea 1-5-2-5 cm.
longa, o-8-i-i cm. lata, setaceo-cuspidata marginibus denticulatis vel leviter
laceratis; bracteolae similes, setaceo-bicuspidatae. Flores pallide lilacini plerum-
que segmentis tribus inferioribus in parte inferiore luteis ; perianthii tubus 2-3 cm.
longus, parte cylindrica 1-3-2-3 cm. longa, superne curvatus gradatim ampliatus
et enguste infundibuliformis ; segmenta obtusa, supremum 2-2*4 cm - longum,
1—1*6 cm. latum breviter et late unguiculatum, lateralia superiora 1-6-2-2 cm.
longa, 0-5-1-1 cm. lata; segmenta inferiora 1*3-2 cm. longa, 4-6 mm. lata,
basin versus leviter attenuata. Stamina clare arcuata; filamenta 1*5-2 cm.
longa; antherae 4 mm. longae. Ovarium 4 mm. longum, 2-3 mm. diam.; stylus
3-3*5 cm. longus ramis filiformibus apice leviter ampliatis.
Corm globose, 2*5-3 cm. diam.; tunics of numerous fine hair-like fibres
extending up in a neck 7-8 cm. long. Stem 28-42 cm. high, simple or with 1
or 2 branches. Leaves 7-9 in the basal rosette, 5-23 cm. long, 0*5-1*4 cm. wide,
acute or subacute, the veins visible but not prominent; 1 cauline leaf near the
middle of the stem, 4-6 cm. long, acuminate. Spikes 2-7-flowered. Bract
1*5-2*5 cm. long, o-8-i- 1 cm. wide, setaceous-cuspidate with denticulate or
slightly lacerate margins, colourless near the base, pale reddish-brown above with
darker red-brown stripes; bracteoles similar, setaceous-bicuspidate. Flowers
mauve or lilac-coloured, usually with the lower half or two-thirds of the
lower lobes yellow; perianth tube 2-3 cm. long, the cylindrical part 1-3-2-3 cm.
long, curved and expanding gradually into the narrowly infundibuliform part;
perianth lobes obtuse, the uppermost 2-2*4 cm. long, i-i*6 cm. wide near the
middle, shortly and broadly unguiculate; upper side lobes 1*6-2*2 cm. long,
0-5-1*1 cm. wide; 3 lower lobes 1-3-2 cm. long, 4-6 mm. wide, slightly
narrowed to the base but not unguiculate. Stamens distinctly arcuate : filaments
A REVISION OF THE GENUS SYNNOTIA 1 47
1-5-2 cm. long; anthers 4 mm. long. Ovary 4 mm. long, 2-3 mm. diam. ; style
3-3-5 cm. long, with filiform branches slightly expanded at the apex.
Type. South Africa, without locality, Roxburgh (not seen).
Clanwilliam Division: Olifants River Valley, 20 miles south of Glanwilliam,
Salter 7495; 15 miles north of Gitrusdal, Lewis (S.A.M. 5791 1); between Clan-
william and Citrusdal, Wilman 850; Kriedouwkrans, between Clanwilliam and
Citrusdal, Barker 5756; Olifants River Valley, Steyn 383.
Flowering season. August to September.
This species is closely allied to S. galeata from which it differs in having
larger flowers with a longer perianth tube, the uppermost lobe comparatively
wider, and the lower lobes not clawed at the base. Baker placed it as a variety
of S. bicolor, with a very brief description — 'Bulb tunics of fine threads ; flower
all lilac-purple'. Unfortunately I have not seen the type but it is almost
certain that the plants described and cited above must be the same. Baker's
description can scarcely apply to S. galeata, in which the flowers, even in dried
material, are not all lilac-purple, and his description of the corm tunics pre-
cludes any likelihood of it being S. Metelerkampiae L. Bol. All the records of
this species are from the same small area in the Olifants River Valley.
5. S. variegata Sweet, Brit. Flow. Gard., t. 150 (1826); Klatt, Linnaea,
xxxii, 750 (1863); Baker, Handbk. Irid., 198 (1892); Fl. Cap., vi, 135 (1896);
S. versicolor Steud., Nom., ed. 2, ii, 614 (1840) ; S. Walthami Hort. Kew, cf. Gard.
Chron. (1881), 1, 370, nomen.
Corm globose or ovoid, 1-2-2 cm. diam.; tunics of hard fibres closely reticulate
above with numerous rather coarse vertical parallel strands in the lower half,
not extending up in a neck but sometimes with one or two old tunics present
above the corm giving the appearance of a neck. Stem 8-40 cm. high, usually
15-20 cm., simple or more often 1 to 3-branched. Leaves 7-8 in the rosette,
2-15 cm. long, 0-5-2 cm. wide, usually 5-10 cm. long and 1-1*5 cm - wide,
obtuse or acute, the veins visible but only the mid-rib slightly prominent;
cauline leaves 1-3, 4-10 cm. long, acute or acuminate. Spike 2-7-flowered,
usually 2-4-flowered. Bract and bracteoles 2-2-5 cm - l° n g> I_I *4 cm - wide,
setaceous-cuspidate with irregularly denticulate margins, usually colourless
in the lower half and pale red-brown above, with a few conspicuous dark red-
brown streaks. Flowers varying in colour from pale mauve and yellow or
cream to deep purple and yellow, the lower lobes sometimes all yellow or
tipped with mauve or sometimes with the upper half or more purple, with
narrow purple and yellow stripes running down into the perianth tube;
perianth tube with the cylindrical part 2-5-4 cm. long, usually 2-5-3 cm -?
geniculate at the top and abruptly dilating into a broad oblique funnel
1-1-5 cm. long and 1-1-5 cm - diam. at the top, the length from the bend in the
perianth tube to apex of uppermost lobe 3-3*5 cm.; uppermost perianth lobe
obovate or oblong-ovate, 2-3 cm. long, 1-2 cm. wide; upper side lobes
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
1-5-2 cm. long, 0-7-1 cm. wide; lower side lobes unguiculate, auriculate above
the claw, 1*3-2 cm. long, 6-8 mm. wide; lowest lobe 1-2-1-5 cm. long, 5-7 mm.
wide. Filaments 1-5-2-2 cm. long; anthers 5-7 mm. long. Ovary 1—5 mm. long,
2-5-3 mm. diam. ; style branches 4-5-6 mm. long, the upper half expanded,
conduplicate and minutely ciliate.
Type. Sweet, Brit. Flow. Gard., t. 150.
Glanwilliam Division: Clanwilliam, Leipoldt, 163, 226; Mader (in Herb.
MacOwan 2138); Olifants River Valley, near Rondegat, Schlechter 5037;
Clanwilliam Barrage, Esterhuysen 5809; Barker 1477; in kloof near Glanwilliam
Barrage, Lewis (S.A.M. 60 11 6); near Vanrhynsdorp, Leipoldt 809; Nardouw
Pass, Barker 7435, 3624; Lewis (S.A.M. 59851); Olifants River Valley, near
Nardouw road, Lewis (S.A.M. 57912); near Langkloof, south of Nardouw
road, Leighton 11 10; 9 miles north of Clanwilliam, Lewis (S.A.M. 60673);
Olifants River Valley Cave, south of Clanwilliam, Compton 22772; 5 miles
south of Clanwilliam, Lewis (S.A.M. 67867).
Flowering season. August to September.
S. variegata Sweet var. Metelerkampiae (L. Bol.) Lewis, comb. nov. S.
Metderkampiae L. Bol., Ann. Bol. Herb., iii, 77 (1923) ; Fl. PL ofS. Afr., hi, t. 98
(1923); Sparaxis luteo-violacea Eckl., Top. Verz., 27 (1827), nomen nudum.
Corm, leaves, inflorescence and bracts as in S. variegata, the corm slightly smaller.
Flowers with shorter and narrower perianth lobes, the length from the bend in
the perianth tube to the top of the uppermost lobe rarely more than 2-5 cm.;
cylindrical part of the perianth tube as in S. variegata, the upper part narrower,
usually 0-5-0-8 cm. diam. at the top, occasionally up to 1 cm.; uppermost
perianth lobe oblong or ovate-oblong, 1-3-2-3 cm. long, 0-7-1-3 cm. wide;
upper side lobes 1-1-5 cm. long, 4-6 mm. wide; lower side lobes 0-9-1-3 cm.
long, 4-6 mm. wide, varying from shortly unguiculate and auriculate above
the claw to subunguiculate and not auriculate; lowest lobe o-8-i-i cm. long,
4-5 mm. wide.
Type. Clanwilliam Division: near Eendekuil, Metelerkamp (B.H. 16039, m
Bolus Herb.).
Clanwilliam Division: near Eendekuil, Metelerkamp (B.H. 16039); Olifants
River, near Brakfontein, Ecklon and £eyher, 120 (76-8) (named Sparaxis luteo-
violacea); Grey's Pass, Steyn 359; Barker 1476; L. Bolus, s. n.; Olifants River
Mtns., behind Warm Bath, Stephens 7091 ; Elands Kloof, Lewis (S.A.M. 57910) ;
Citrusdal, Barker 3761; 15 miles north of Gitrusdal, Lewis (S.A.M. 67865);
Hall 541; 20 miles north of Gitrusdal, Lewis (S.A.M. 67866); near Pakhuis
Leipoldt (B.H. 21277); Pakhuis Pass, Gillett 4062; Salter 3641, 7505; Lewis
(S.A.M. 59849); Compton 20941; Barker 4700; Graafwater, Compton 24220;
sandveld between Grey's Pass and Graafwater, Leipoldt 3583; between Berg
Valley and Clanwilliam, Lewis (S.A.M. 59848) ; Lamberts Hoek Berg, Maguire
415; Barker 671 1; Cedarberg, c. 2,000 ft., Bodkin, s. n.; Algeria Forest Station,
Barnes (B.H. 19451); Algeria Forest Reserve, rocky slope, 1,000 ft., Story 2938.
Flowering season. August to September.
A REVISION OF THE GENUS SYNNOTIA 1 49
Since the publication of S. Metelerkampiae a little over thirty years ago, a
great deal more material has been collected which has proved it to be much
more closely allied to S. variegata than was apparent from the limited amount
of material available at that time. After examining a large number of speci-
mens I have come to the conclusion that it is too closely linked with S. variegata
through intermediate forms to be treated as a separate species and have
accordingly placed it here as a variety.
S. variegata has been recorded from various localities extending from about
five miles south of Clanwilliam through the Olifants River Valley to the top
of the Nardouw Pass, the flowers varying in colour according to the locality,
from purple marked with yellow south of and in the vicinity of Clanwilliam,
to pale mauve and yellow from Clanwilliam northwards. The perianth lobes
are nearly always longer and considerably broader than in the variety,
especially those of the pale colour form from the northern limits, and the
funnel-shaped part of the perianth tube is broader and frequently has a rounded
and somewhat pouch-like curve on the lower side, usually more conspicuous
in the pale colour form.
The type of S. Metelerkampiae represents the extreme southern form of the
variety, but a larger form from the Olifants River Valley, about ten miles south
of Clanwilliam, clearly links this up with the southern form of S. variegata. The
red colouring on the 3 lower lobes in the type is unusual and in the majority of
specimens seen the flowers are either dark purple throughout or have small
spathulate yellow markings near the base of the lower side lobes. _
At low altitudes the plants of the variety are as a rule between 18 and 25 cm.
high but at altitudes from about 1,500 to 2,000 ft., i.e. from Lamberts Hoek
Berg, Pakhuis Pass and slopes of the Cedarberg, the plants are mostly shorter
and many are distinctly stunted with stems in some cases not more than 1 or
2 cm. high. Specimens collected by Leipoldt in 1941 (no. 3872) are recorded as
coming from the Calvinia Division, between Nieuwoudtville and Oorlogs
Kloof, but it is doubtful if this locality is correct.
Species Excluded
S. stenophylla Baker, Bull. Herb. Boiss., Ser. II, 1, 865 (1901).
The type, Ecklon and £eyher 118, is in the Herbarium of the Botanical
Museum, Zurich, and there is an isotype in the South African Museum
Herbarium. By courtesy of Prof. Daniker, who kindly sent the type to me for
examination, it has been possible to clear up the uncertainty regarding the
identity of this species, which proves to be a mixture of Sparaxis and Synnotia.
Baker made an unfortunate mistake in describing this species, partly due to
an error in mounting, and described the plants of a Sparaxis and the flower of a
Synnotia. In a small envelope attached to the type sheet there are two separate
and entirely different flowers. One of these, without bracts, is a flower of
I50 ANNALS OF THE SOUTH AFRICAN MUSEUM
Synnotia variegata var. Metelerkampiae, which Baker erroneously described as
belonging to the plants on the sheet. He evidently overlooked the remains of
a small yellowish flower attached to one of the specimens on the sheet, also the
second flower in the envelope, which is complete with bracts and is the same
as the one on the sheet, though in better condition. On the isotype in the South
African Museum Herbarium there are several flowers, all yellow in colour
and in a good state of preservation. This is a species of Sparaxis and there is no
Synnotia plant on the type or isotype sheets. The odd Synnotia flower placed
with the type specimens was obviously put there by mistake and most probably
came from Ecklon and £eyher no. 120, which is Synnotia variegata var.
Metelerkampiae.
A REVISION OF THE GENUS SYNNOTIA
J 5
Map showing distribution of Synnotia species.
1 . S. parviflora.
2. S. villosa.
3. S. galeata.
4. S. Roxburghii.
5. S. variegata.
6. S. variegata var. Metelerkampiae.
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