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A Synopsis of Woody Portulacaceae in Madagascar

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A Synopsis of Woody Portulacaceae in Madagascar A synopsis of woody Portulacaceae in Madagascar Urs EGGLI Stádtische Sukkulenten-Sammlung, Mythenquai 88, CH-8002 Zurich, Switzerland. ABSTRACT The relationships of the endemic woody Madagascan genera Talinella Baillon and Dendroportulaca (gen. nov.) are discussed. Talinella (5 species) is unique in the family for its juicy-mucilaginous berry-like fruits, while Dendroportulaca (monotypic) has unique spicate inflorescences but shares operculate fruits with the herbaceous genus Portulaca. Talinella and Dendroportulaca, which share similar smooth black and apparently primitive seeds, are probably primitive within the phylogeny of the family, and their relationships to the African woody genera Ceraria and Calyptrotheca KEYWORDS (Portulacaceae) as well as to Basellaceae are discussed. The genus Talinella is Portulacaceae, monographically revised and a key to its 5 species is presented. T. microphyl- Talinella, Dendroportulaca, la and T. pachypoda are described as new. Dendroportulaca is monotypic Madagascar. with D. mirabilis also newly described. All taxa are illustrated. RÉSUMÉ Les affinités des genres malgaches, endémiques et ligneux, Talinella Baill. et Dendroportulaca (gen. nov.) sont discutées. Au sein des Portulacaceae Talinella (5 espèces) est unique par ses fruits bacciformes juteux-mucilagi- neux, et Dendroportulaca est unique par des inflorescences en épis tout en présentant des fruits operculés comme dans le genre herbacé Portulaca. Talinella et Dendroportulaca, qui possèdent en commun des graines noires, lisses et apparemment primitives, ont probablement une position primirive dans la phylogénie de la famille et leurs affinités avec les genres ligneux afri­ cains Ceraria et Calyptrotheca (Portulacaceae), de même qu'avec les Basellaceae, sont discutées. La révision de Talinella est présenrée sous une MOTS CLÉS forme monographique avec une clé de détermination des 5 espèces. Deux Portulacaceae, nouvelles espèces, Talinella microphylla et T. pachypoda sont décrites. Talinella, Dendroportulaca est monotypique et l'espèce D. mirabilis est également Dendroportulaca, Madagascar. décrite pour la première fois. Tous les taxons sont illustrés. sér. 3 • 1997 • 19 (1) 45 Eggli U. INTRODUCTION of Fort Dauphin, and in 1915, DANGUY descri­ bed T grevei from near Morandava, comparing The recent interest in caudiciform (term intro­ it with T. dauphinensis. duced by CD. ROWLEY in 1948 for plants with Half a century later, LEANDRI (1962) published succulent basal stotage organ and ± mesophytic, the new monotypic genus Sabouraea which he often annual, aerial stems; ROWLEY 1987) plants associated, though with a question mark, with among collectors of succulents and other xero- the Flacouttiaceae. Only three years later, phytes has brought to light many interesting taxa LEANDRI (1965) recognized the synonymy of from Madagascar, predominantly of genera of Sabouraea with Talinella, and reported that the Cucurbitaceae (e.g. Trochomeriopsis, Xerosicyos) parietal placentation indicated in the illustrations and Passifloraceae {Adenia), but also occasionally which accompanied the protologue of Sabouraea from other systematic groups. must be an error of observation. A careful exami­ Such an introduction was also the starting nation of Sabouraea sarmentosa now reveals that point for the investigations which led to the pre­ it is conspecific with Talinella grevei. Talinella sent paper: Some years back, the author received does not seem to have been the subject of further an unidentified caudiciform from the extreme N studies, with the exception of the superficial and of Madagascar. It was soon clear that this plant uninformative study of NYANANYO (1986) who was a member of the genus Talinella, but concluded on the basis of stomatal patterns that attempts to identify it as to species were unsuc­ the genus properly belongs to the Portulacaceae. cessful and showed that the genus was in need of Since the species of Talinella are virtually un­ revision. known in cultivation (except some collections cultivated in the Heidelberg Botanic Garden for many years, and the more recently introduced HISTORICAL SUMMARY material which was the reason to initiate this study), it is also absent from contemporary suc­ Members of Portulacaceae are not conspicuous culent plant literature (e.g. RAUH 1995)—a amongst the xerophytic and succulent vegetation notable fact since its taxa appear to be quite of Madagascar (e.g. not mentioned in PvAUH widespread in the xetophytic vegetation of SW 1995). Casual examination of herbarium hol­ Madagascar. The synoptical revision that follows dings shows that apart from Talinella (3 species is mainly based on herbarium specimens at K, described) there are a handful of species of MO, P and Z, as well as on ample living matetial Portulaca and the occasional specimen of of Talinella pachypoda. Talinum, the latter no doubt representing neo- Checking the folders with undetermined phytic weeds. Portulacaceae for material of Talinella revealed Talinella was described by FJAILLON (1886a, the presence of a second woody-shrubby genus 1886b) with the single species T. boiviniana, of similar appearance in Madagascar. Despite the whose geographic origin within Madagascar was paucity of material available, it will subsequently not mentioned. Its primary distinctive charactet be described as Dendroportulaca, to make the according to FJAILLON is the ovary, which is divi­ name available for a forthcoming treatment of ded into two locules—a character which is the family for the Flore de Madagascar. unique in the family and which makes the syste­ matic placement of the genus difficult (PAX & HOFFMANN 1934: 261; CAROLIN 1993). FRANZ GENERIC RELATIONSHIPS (1908: 19) even went so far as to exclude the genus from the Portulacaceae, speculating about Relationships amongst the genera of possible affinities to the Aizoaceae on account of Portulacaceae are still insufficiently known the bilocular ovary. (HERSHKOVITZ 1993; HERSHKOVTTZ in ms.) and A second species, T. dauphinensis, was descri­ this is equally true for the whole group of fami­ bed by SCOTT ELLIOT in 1891 from the vicinity lies, including Didiereaceae, Basellaceae and 46 ADANSONIA, ser. 3 • 1997 • 19(1) Woody Portulacaceae in Madagascar Cactaceae (HERSHKOVITZ in litt.). Since no spe­ date it. Incidentally, NYANANYO (1990) placed cific ancestral taxa (outgroups) can be recogni­ Talinella in tribe Talineae, but the fruit characte­ zed, character interpretation and cladistic studies ristics are incongruent with his definition of the are at present difficult to carry out. Both PAX & tribe. HOFFMANN (1934) and CAROLIN (1993) placed The position of Talinella was also discussed by Talinella in the family Portulacaceae, but as a HERSHKOVITZ (1993). He places the genus in his genus of very uncertain position. While its gene­ "American/African group of the portulacaceous ral habit—apart from its sarmentóse growth—is alliance", suggesting closest affinities with rather suggestive of some taxa of Talinum (e.g. Didiereaceae and Ceraria and Portulacaria. T. portulacifolium (Forssk.) Asch. ex Schweinf.) A further character of interest is the occurrence and is responsible for the generic name, several of unisexual flowers in Talinella, probably cou­ flower and fruit characters are notable: The sep­ pled with completely dioecious individuals at tate ovary was already mentioned as an exceptio­ least in some cases. The available herbarium nal character by BAILLON (1886a), and its material does not allow firm conclusions, but it presence was recently confirmed by HERSH­ appears that completely unisexual inflorescences KOVITZ (pers. comm.). HERSHKOVITZ found that are rather the rule than the exception, and that material probably representing T. grevei (speci­ apparently bisexual flowers could well be functio­ men not recorded) had a bilocular ovary, each nally unisexual, with either underdeveloped (or locule with a single ovule basally attached to the lacking) anthers or underdeveloped ovaries. All septum. In T. boiviniana {Humbert 18878, P), plants of T. pachypoda investigated are dioecious, HERSHKOVITZ found an incompletely 4-locular and complete or partial dioecy is another rare ovary, i.e. a complete primary septum as in T. character in the family, associated with woody grevei, with 2 additional incompletely formed habit in Ceraria and Portulacaria. septae in the form of peg-like protrusions direc­ A comparison with other genera of ted from the ovary wall towards the centre. This Portulacaceae will have to include woody mem­ is in accordance with a notes made by BAILLON bers with similar growth. These are Ceraria and (1886a) on the basis of information supplied by Calyptrotheca, both from the African continent. the collector BoiVTN that the fruit was 4-locular Calyptrotheca (2 species in E Africa): This genus with 1 seed per locule. is similar in its woody habit (though not sarmen­ Unfortunately, BoiVIN made no notes about tóse), semi-succulent deciduous leaves, inflores­ the fruits of T. boiviniana, but remarks on seve­ cence position and architecture (esp. the ral herbarium specimens of T. grevei and obser­ abbreviated contracted part-inflorescences with ± vations of living material of T. pachypoda show dichasial branching and the small bracts), basally that the fruits exhibit another exceptional charac­ hairy filaments, similar pollen (NYANANYO 1986: ter for the family, as they ripen into juicy mucila­ tab. 62; NYANANYO 1989), few-seeded fruits and ginous berries. In T. pachypoda, these berries are smooth
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