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Convincing Habit, Viz. Climbing

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Convincing Habit, Viz. Climbing The systematic position of tribe Paropsieae, in particular the genus Ancistrothyrsus, and a key to the genera of Passifloraceae W.J.J.O. de Wilde Summary The members of tribe Passifloreae of the Passifloraceae are assumed to be originally all tendril-climbers. have and the ramification They essentially axillary cymose inflorescences, vegetative occurs always through the bud. In tribe and Flacourtiaceae shrubs the accessory Paropsieae (mostly or trees, no tendril-climbers) inflorescences are axillary, most likely essentiallyracemose, and the vegetative ramification is mostly through the axillary bud. The tendril-climbing neotropical genus Ancistrothyrsus appeared to belong to Passifloreae. the tribe remains and Though Paropsieae to occupy an intermediate position between Passifloraceae Flacourtiaceae, they can best be classified with the Passifloraceae. A new key is proposed for the distinction of both families and the genera of Passifloraceae. the of revision of the Adenia, and the treatment of the During preparation my genus Passifloraceae for the Flora Malesiana, I became of course interested in the systematics of the of the tribe several the family, amongst others in position Paropsieae on which subject that this papers were recently published. Almost all authors agree tribe occupies a more or less transitional place between Flacourtiaceae and Passifloraceae. These two families are certainly more closely allied mutually than to others and show consequently also phytochemical similarities (Hegnauer, 1969). The tribe first also was recognized by De Candolle (1828) in Passifloraceae; e.g. Perrier de la Bathie (1945) and Fernandes (1958) included it in that family. By most British authors, from Bentham & Flooker (1867) onwards to Hutchinson (1967), the genera it contains without the of are accommodated in Passifloraceae but recognition a supra -generic taxon. and For not very convincing anatomical reasons the different habit, viz. arboreous the tribe transferred versus climbing, was to the Flacourtiaceae by Harms (1893), an modern opinion followed until time by various German authors, e.g. Engler (1921), Gilg (1923), Melchior in Engler's Syllabus (1964), and Sleumer (1954). Recent anatomical and induced Sleumer palynological investigations, however, (1970) to refer the genus Paropsia to Passifloraceae. Since the of it has treatment Paropsieae by Gilg (1925) appeared that the genus Soyauxia from the with should be excluded tribe, whereas Hounea proved synonymous Paropsia. Two viz. liana Peru newParopsiaceous genera have been added, Ancistrothyrsus, a from East Brazil and West (Harms, 1931, 1932) and Viridivia, a shrubby tree closely related to Paropsia, from Rhodesia (Hemsley & Verdcourt, 1956). the At present tribe Paropsieae is composed of the following genera: Paropsia, Andro- small siphonia, Viridivia, Paropsiopsis, Smeathmannia, and Barteria, all shrubs or trees from the Old World, and Ancistrothyrsus, a liana from South America. 100 BLUMEA VOL. XIX, No. i, 1971 When included the tribe the in Flacourtiaceae, is distinguished mainly by presence of a corona. Within Passifloraceae it is remarkable that the Paropsieae, with the exception of South the climbing American Ancistrothyrsus, are deviating by their arboreous habit; the tribe are all tendril-climbers with a few the genera constituting Passifloreae herbaceous, Adenia erect derivatives in and Tryphostemma and a few shrub-like outliers in the large the related genus Passiflora and genus Dilkea. Malesian been During myinvestigations in Adeniaand Passifloraceae I have impressed by the uniform modeoframificationin Passifloraceae, namely through thesecond, serial, acces- bud. Harms in Nat. Pfl. sory Already (1893, 1925) wrote (family diagnosis Fam., 1925, seriale p. 471): 'oberhalb der Ranke oder des Bliitenstandes eine Beiknospe, die zu einem Laubspross oder durch Reduktion der Blatter zu einem Bliitenstand auswachsen kann'. Searching for characteristics for the distinction of the tribe Paropsieae within the Passi- floraceae better than 'shrubs or trees without tendrils' versus 'climbers, mostly provided tendrils' I checked the of and with presence the serial accessory buds the corresponding of the mode of (vegetative) ramification in all the genera Passifloreae: Passiflora (inch Tetrastylus), Dilkea, Mitostemma, Hollrungia, Tetrapathaea, Adenia, Tryphostemma, Cros- and sostemma (inch Schlechterina), Deidamia (inch Efulensia), the Paropsieae, and a large number of Flacourtiaceae. This revealed the following: all 1) In Passifloraceae (Passifloreae & Paropsieae) an axillary and an accessory bud are Flacourtiaceae bud always present; in an accessory is either present (e.g. in Lindackeria, absent. Ryparosa, Hydnocarpus, many Casearias) or all is into 2) In Passifloreae the axillary bud either abortive, or developed a cymose sterile inflorescence (either tendril-bearing or not) or into a tendril; the second, serial, the accessory bud always provides for vegetative ramification, whereas in certain A. species (e.g. Passiflora racemosa, Tetrastylis ovalis, Adenia racemosa, globosa, Dilkea sp.) this raceme-like i.e. shoot appears as an inflorescence, an inflorescences-bearing or The short-shoot. ultimate 'true' inflorescences, which are essentially cymose, never from the bud. a—d. develop accessory Fig. 1 the the bud flower- 3) In Paropsieae axillary develops into a or inflorescences-bearing also for the ramification. The bud either abor- shoot; it provides vegetative accessory is tive or rarely developed into a normal branch. In the flower- or inflorescences shoots the flowers inflorescences situated —h -bearing or apparently are axillary. Fig. I e 4) In the Flacourtiaceae the ramification generally resembles that in Paropsieae. The bud for the it axillary usually provides vegetative ramification, or may produce a racemose inflorescence (raceme-like or thyrsoid e.g. in Homalium, Lindackeria, Casearia, Ryparosa, Hydnocarpus, and others), or a flower-bearing short-shoot, sometimes reduced to fas- African cicles or even to a single flower (e.g. the Australian Streptothamnus). In the South genus Kiggelaria the vegetative ramification seems to occur always through the accessory the bud, but axillary inflorescences are racemose, i.e. flower-bearing short-shoots. Fig. I e, k—p. The difference in mode of ramification between the Passifloreae on the one hand, and the Paropsieae and Flacourtiaceae on the other, leads me to the assumption that the inflores- in the the uni- cences two latter groups are likely essentially racemose. Whether simple or few-flowered ultimate inflorescences, as found e.g. in Paropsia or various Flacourtiaceae (e.g. Casearia, Hydnocarpus, etc.), are essentially reduced racemes or cymes remains, of course is difficult to decide. If my observations and considerations on the mode of ramification are correct (which this rather needs additional checking on several genera of Flacourtiaceae), would mean a fundamental difference betweenPassifloreae and Paropsieae (together with Flacourtiaceae), W. J. J. O. db Wilde: The systematic position of the Paropsieae 101 Fig. 1. Mode of vegetativeramification and various inflorescences in a—d. Passifloreae, e—h. Paropsieae, Essence in and e, k—p. some Flacourtiaceae; all diagrammatic. Accessory bud solid black, a. of ramification A. Passifloreae; b. Most Passifloras and Adenias; c. Passiflora racemosa, Tetrastylis ovalis, Adenia racemosa, P. Essence venenata; d. Passiflora arborea, spicata, Dilkea, Adenia globosa, A. fasciculata; e. of ramification in and f. Paropsieae Flacourtiaceae; Paropsia, Paropsiopsis, Smeathmannia; g. Paropsia guineensis, Androsiphonia; h. k—n. Various of Bacteria; genera Flacourtiaceae, e.g. Homalium, Lindackeria, Ryparosa, Scolopia p.p., dichasial inflorescences in Laetia Laetia. Hydnocarpus, etc.; o. Phyllobotrieae; p. Axillary sect. in addition the rather character used till of to only arbitrary up now 'climbing' versus This would 'arboreous'. carry even more weight if a third differential character would be 'inflorescences essentially cymose' against 'inflorescences predominantly racemose'. To these to the few of test assumptions it appeared interesting investigate cases excep- in tional habit both groups. In of the P. arborea and Passifloreae some species large genusPassiflora are arboreous, e.g. the consists P. spicata; genus Dilkea mainly of non-climbing shrubs of small trees; and in Adenia and Tryphostemma some erect non-climbing herbaceous species occur. It appeared that the mode of ramification in all these species is essentially the same as in of the Dilkea the 'normal' climbing Passifloras, as explained above. Checking genus rather hand. the gave some difficulties because of the poor material at Here situation in the ramificationis usually obscured by the fact that the true axillary bud is suppressed; in a few cases, however, an abortive tendril between leaf scar and inflorescence (i.e. a flower- is other bearing short-shoot) revealed that thesituation exactly as in Passifloreae. Moreover, Dilkea is closely related to Mitostemma, in which the situation is quite the same as in Passiflora. The few scandent Flacourtiaceae, i.e. the Australian Streptothamnus and Berberidopsis No. 102 BLUMEA VOL. XIX, I, 1971 a—b. Fig. 2. Two remarkably different flowers ofAncistrothyrsus tessmannii Harms. Longitudinal section and separate androgynoecium of a mature flower from Ducke 24385, anthers non-apiculate, x 3. c.—d. Ditto, from Ducke 35681, a small-flowered specimen; note the longly apiculate anthers, x
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