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Full text of "Annals of the South African Museum = Annale van die Suid-Afrikaanse Museum"

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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XXX. i93i-*935 Zoology 

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XXXVI. 1942-1948 Zoology 

XXXVII. 1947- Archaeology . (Parti) 

XXXVIII. 1950 Zoology 

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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 

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Univ. CXXVII, 47-8 (1939). 
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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). 



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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 



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SOUTH AFRICAN MUSEUM 

VOLUME XL 

PART IV, containing: — 

5. A revision of the Genus Synnotia. By G. J. Lewis, B.A., Ph.D. 





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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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XXXIX. 1952 Zoology 2 14 6 

XL. 1952- Botany .. Parti, is.; 2, ijs.6d.; 3,5^. Part 4 030 

XLI. 1952-1955 Zoology . . .. 400 

XLII. 1953- Palaeontology .. .. Part 1, 12s. 6d. Part 2 o 15 o 

XLIII. 1955- Zoology Part 1 100 



Copies may be obtained from — 

The LIBRARIAN, South African Museum, Cape Town, 

except the Geological and Palaeontological parts, which are obtainable from the 
Government Printer. Pretoria. 



J 






SMITHSONIAN INSTITUTION LIBRARIES 




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