Bothalia 11, 1 & 2: 191-201 (1973) Notes on the genus Cussonia in South Africa R. G. S T R E Y * ABSTRACT Information gathered during studies o f the genus Cussonia under field conditions is presented. The development o f the inflorescences, and branch and leaf initiation, is given special attention. The subdivision o f the genus Cussonia is discussed and the following new subgeneric taxa are created: Subgenus Cussonia (type species: C. spicata Thunb.) with two sections: Sect. Cussonia and Sect. Capitata Strey; Subgenus Paniculata Strey (type species: C. paniculata Eckl. & Zeyh.); Subgenus Protocussonia (type species: C. natalensis Sond.). It is pointed out that the names C. kraussii Hochst. and C. chartacea Schinz have been misapplied to two undescribed species, and are synonyms o f C. spicata Thunb. and Schefflera umbellifera (Sond.) Baill., respectively. Four new species are described viz. C. arenicola Strey, C. zuluensis Strey, C. nicholsonii Strey and C. sphaerocephala Strey. INTRODUCTION by the bract-covered apices from which the in- In the course o f studies o f the genus Cussonia a florescence-branches develop. The development of number o f interesting facts emerged, especially from the inflorescence o f two o f the species studied differs the field studies that were undertaken. Several of fundamentally from the pattern displayed by the these features, particularly the development o f the others. In most species the inflorescence bud initials inflorescence, the mode o f flowering and the renewal are laid down in the bract-covered apices of some of vegetative growth after flowering, have apparently o f the branches towards the end o f the growing not been recorded before and are therefore discussed season and develop from these usually in the following in the following pages. It was also discovered that spring. The situation in Cussonia spicata and C. two names had been misapplied and that the species sphaerocephala is quite different. Instead of producing masquerading under these names were new and inflorescences, an “ umbel” of fairly slender leafy required to be described. In addition, two new species branches is produced at the apex of a trunk during were identified and are described here. I am indebted the first season. Each branch has a knob-like thick­ to Dr D. J. B. Killick for the Latin descriptions. ening covered with bracts, at the apex. Only during The majority of specimens studied are deposited in the follow ing season are inflorescences produced at the National Herbarium, Pretoria, and Natal Her­ the apices o f these branches. Since the leafy branches barium, Durban. Those seen in other herbaria are formed during the first season lose their year-old specially indicated. leaves, as soon as the inflorescences start developing, the final structure resembles a compound “ umbel” at the end o f the second season. These pseudo­ FIELD STUDIES inflorescences are therefore biennial structures in The development o f the inflorescence contrast to the inflorescences produced by other species which develop in one year. In all species o f Cussonia the structures bearing the flowers are borne in clusters at the apices o f The flowers and fruits branches or in some cases even at the apices o f The fully-developed flowerbuds open up in irregular fairly thick trunks. These clusters usually consist o f sequence on almost any parts o f the spikes or racemes, a few to many peduncled spikes, racemes or even at the base or the apex or at the middle, and in panicles originating in the axils o f the apical bracts addition one side o f the spike may be in full flower which are numerous and in dormant branches com­ when no flowers have opened on the other side. pletely cover the apices. A t the beginning o f a new Flowers at various stages of development are there­ season when the trees start sprouting it is soon evident fore found irregularly grouped in the inflorescences. that branch apices either produce a new flush o f The open flowers are star-shaped with the petals and leaves or somewhat more slowly develop inflorescen­ stamens stiffly spreading. The petals and stamens ces. Leaves and inflorescences are seldom, if ever, are very easily detached and are never preserved simultaneously produced at the apex o f the same in situ in dried specimens. branch. Whereas branches producing inflorescences Newly opened flowers are greenish-cream to may be leafy, these leaves are almost invariably butter-yellow in colour, the top o f the ovary green, produced during the previous season and are usually the stigmas pale and erect and the anthers yellow. shed before the inflorescences are mature. Mature As the flower ages the petals and stamens drop, the and particularly fruiting inflorescences are therefore top o f the ovary becomes yellow and exudes a shiny, borne at the apices o f bare branches. In all the species sticky fluid which attracts insects, the stigmas become studied, the clusters o f pedunculate flower-bearing darker and the styles recurve. It is probably at this branches together form an apical compound in­ stage that pollination is effected by the various types florescence, since no normal leaves are produced o f insects found on the flowers, including bees, flies ‘ Botanical Research Unit, Botanic Gardens Road, Durban. and beetles. Not all ovaries are fertilized. The ones 192 NOTES ON THE GENUS CUSSONIA IN SOUTH AFRICA that are fertile increase in size and develop a fleshy Leaf-form and development purple exocarp attractive to birds. Each fruit con­ The leaves, which in all species are clustered at tains one to two seeds. Unfertilized and parasitized the ends o f branches, are, in most species, produced ovaries may develop but remain dry and hollow in flushes. The young leaves mature and usually structures. Mature inflorescences are attacked by remain on the tree for a full year. Old leaves lower insects and the stem-apex in particular seems sus­ on the stems are gradually cast o ff after a new flush ceptible. When the stem-apex has decayed, the whole o f leaves has appeared so that the clusters are always inflorescence breaks off and drops to the ground. situated apically. Usually only one flush o f leaves per year is produced. Renewal of leaves, as also the The formation o f new branches production of inflorescences, is not only seasonally The apices o f the stems which have fruited and determined and may vary from tree to tree. have subsequently lost the inflorescences, are now In shape the leaves vary from simple and deeply usually more or less truncate and dry out, or develop lobed to singly or doubly compound. The leaflets a certain amount o f callus-tissue. These truncated o f the compound leaf are arranged digitately (like the branches and trunks eventually produce side-shoots spokes o f a wheel) round the apex o f the peduncle. which continue the growth o f the branch. It appears They spread in one plane and at an angle to the that side-shoots are seldom formed on actively growing peduncle forming a fan-like structure, sub-circular vegetative branches and trunks in the sparingly in outline. The individual leaflets are vertebrate, branched species such as C. spicata and C. sphaero- a term first used by Mirbel (1815, p. 655) for leaves cephala, but that usually one, rarely more, side- o f Cussonia spicata. These leaflets resemble pinnae branches are formed after the growth o f a trunk or with the rachillas very broadly winged. The leaf branch has been temporarily halted by production therefore is a doubly compound structure with the o f an inflorescence or by injury. In species which first division digitately compound the second verte­ by nature have a more branched habit, such as brate or rarely doubly vertebrate. The leaflets are C. natalensis, several shoots originate from dormant arranged in pairs, lower pairs being the least de­ buds after the inflorescences have been discarded. veloped and the single terminal leaflet showing the pin n rhachis lobe VERTEBRATE LEAFLET lowest leaflet Fra. I.— Semi-diagrammatic drawing of a leaf of Cussonia spicata to illustrate the terminology used in this paper. R. G. STREY 193 most elaborate development. The vertebrate leaflets A second distinct group is represented by only have 1-5 articulations, characterized by turgid green one species, namely, C. paniculata Eckl. & Zeyh. papillate scales on the lower suface. These scales which may be distinguished from all other species may represent reduced bracts. A t these articulations by the paniculate branches of the inflorescence and the pinnules are attached. The articulations may the elongate simple, shallowly to deeply lobed leaflets be devoid o f pinnules, or bear two, four or six pinnules which are never vertebrate. arranged in a whorl, the numbers increasing pro­ The third group is composed of species which gressively upwards. The terminal pinnules o f the have palmatifid. palmatisect or digitately com­ vertebrae usually repeat the form of the lowest pound leaves. There seems to be a somewhat gradual leaflet, if this is a simple structure. Because o f the transition from palmatilobed— palmatifid— palmati­ size o f the compound leaves, only single leaflets are sect— digitately compound leaves (see also Aitken. usually represented in herbaria and the leaf-structure 1923, p. 58), and a further division o f the species is usually not fully understood. In Fig. 1 a complete into two groups, one possessing simple lobed leaves leaf of C. spicata is semi-diagrammatically represented. (e.g. C. natalensis) and a group possessing compound This figure also serves as a reference to the terminology palmatisect or digitate leaves was contemplated used in this paper.