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Endemic Families of Madagascar. V. a Synoptic Revision of Eremolaena, Pentachlaena and Perrierodendron (Sarcolaenaceae)

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Endemic Families of Madagascar. V. a Synoptic Revision of Eremolaena, Pentachlaena and Perrierodendron (Sarcolaenaceae) Endemic families of Madagascar. V. A synoptic revision of Eremolaena, Pentachlaena and Perrierodendron (Sarcolaenaceae) Porter P. LOWRY II Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, U.S.A. [email protected] Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle, 16 rue Buffon, 75005 Paris, France. [email protected] Thomas HAEVERMANS Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle, 16 rue Buffon, 75005 Paris, France. [email protected] Jean-Noël LABAT Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle, 16 rue Buffon, 75005 Paris, France. [email protected] George E. SCHATZ Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, U.S.A. [email protected] Jean-François LEROY† Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle, 16 rue Buffon, 75005 Paris, France. Anne-Elizabeth WOLF Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle, 16 rue Buffon, 75005 Paris, France. [email protected] ADANSONIA, sér. 3 • 2000 • 22 (1) : 11-31 © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. 11 Lowry II P.P. et al. ABSTRACT As part of an assessment of the vascular plant families endemic to Madagascar and the Comoro Islands, a synoptic revision is presented of three genera of Sarcolaenaceae: Eremolaena Baill. (2 spp.), Pentachlaena H. Perrier (3 spp.), and Perrierodendron Cavaco (5 spp.). Molecular sequence data place Sarcolaenaceae as the sister group of Dipterocarpaceae within an expanded Malvales. The three genera studied appear to be closely related and probably form a monophyletic assemblage, sharing several diagnostic features (e.g., free stipules and inflorescence bracts, an involucre reduced at anthesis but expand- ing somewhat in fruit in certain species, and seeds with little or no endosperm KEY WORDS Sarcolaenaceae, and a smooth integument); features separating the genera include sepal size, Eremolaena, the number of carpels, and fruit dehiscence. Five species are described as new, Pentachlaena, including one Pentachlaena (P. betamponensis) and four Perrierodendron (P. Perrierodendron, Madagascar, capuronii, P. occidentalis, P. quartzitorum and P. rodoense). Keys to the genera endemism. and species are provided in English and French. RÉSUMÉ Familles endémiques de Madagascar. V. Révision synoptique des genres Eremolaena, Pentachlaena, et Perrierodendron (Sarcolaenaceae). Dans le cadre de l’évaluation des familles de plantes vasculaires endémiques de Madagascar et des Comores, la révision synoptique de trois genres apparte- nant aux Sarcolaenaceae est présentée : Eremolaena Baill. (2 spp.), Pentachlaena H. Perrier (3 spp.), et Perrierodendron Cavaco (5 spp.). Des ana- lyses moléculaires montrent que les Sarcolaenaceae sont le groupe frère des Dipterocarpaceae, au sein des Malvales élargies. Les trois genres étudiés sem- blent être étroitement apparentés entre eux et forment vraisemblablement un groupe monophylétique, partageant plusieurs caractères diagnostiques (ex. : des stipules libres ainsi que les bractées inflorescentielles ; un involucre réduit à l’anthèse mais se développant quelque peu lors de la fructification chez cer- taines espèces, des graines avec peu ou pas d’endosperme et un tégument lisse) ; les caractères distinctifs des genres sont, entre autres, la taille des MOTS CLÉS Sarcolaenaceae, sépales, le nombre de carpelles et la déhiscence du fruit. Cinq espèces nou- Eremolaena, velles sont décrites, un Pentachlaena (P. betamponensis) et quatre Perriero- Pentachlaena, dendron (P. capuronii, P. occidentalis, P. quartzitorum et P. rodoense). Des clés Perrierodendron, Madagascar, de détermination des genres et des espèces sont établies en anglais et en endémisme. français. INTRODUCTION vation status of the approximately 100 species concerned will be assessed (SCHATZ et al. 1998, This is the fifth in our series of synoptic revi- 1999a,b, in press; LOWRY et al. 1999). We have sions of genera belonging to Madagascar’s eight examined all the available material of three endemic plant families, each of which is being closely related genera of Sarcolaenaceae prepared to provide a modern framework for (Eremolaena Baill., Pentachlaena H. Perrier, and compiling a Red Data Book in which the conser- Perrierodendron Cavaco) at the major herbaria 12 ADANSONIA, sér. 3 • 2000 • 22 (1) Eremolaena, Pentachlaena and Perrierodendron (Sarcolaenaceae) with important holdings of Malagasy plants (K, cladistic analysis of the family using morphologi- MO, P, TAN and TEF), and have reviewed the cal features (HAEVERMANS 1999). A recent study circumscription of species and infraspecific taxa, of pollen morphology among Sarcolaenaceae as presented by CAVACO (1951, 1952a,b) and (NILSSON & RANDRIANASOLO 1999) indicates updated by CAPURON (1973). that the tetrads of all but one species in these CAVACO, in his treatment of Chlaenaceae (= three genera are similar in structure, with promi- Sarcolaenaceae) for the Flore de Madagascar nent apertural ridges comprising disk-like lumps (1952b; see also 1952a), recognized two species or claw-like elements, a finding that is consistent of Eremolaena and a single species each of with their hypothesized close relationship. Pentachlaena and Perrierodendron. Two decades However, the pollen of the single sample of E. later, CAPURON (1973) described a second species rotundifolia studied is distinctly different, com- of Pentachlaena. Shortly thereafter, J.-F. LEROY prising single grains with a reticuloid-rugulate, began drafting a paper describing four new columellar exine – an observation that would be species of Perrierodendron, which unfortunately worth confirming in additional material. remained unpublished at the time of his death Notwithstanding this possible palynological early last year. A careful study of the herbarium anomaly, however, the three genera appear to be material of this genus has confirmed that all four closely related, and can be separated from one taxa are sufficiently distinct to warrant recogni- another using several features, including sepal tion at the species level. Analysis of the specimens size, the number of carpels, and fruit dehiscence. of Pentachlaena has also revealed a third unde- Examination of the available material of scribed species in that genus. Eremolaena, Pentachlaena, and Perrierodendron Based on recent molecular sequence data, has led us to propose the following revised taxon- Sarcolaenaceae and Dipterocarpaceae comprise omy, in which ten species are recognized, five of sister taxa in a group that also includes Cistaceae which are described as new. The epithets chosen and Muntingia L. within an expanded Malvales for two of the new species (Perrierodendron (ALVERSON et al. 1998; BAYER et al. 1999). Fossil quartzitorum and P. rodoense) were taken from pollen recorded from the Miocene of South annotations made by CAPURON and LEROY, Africa clearly belongs to Sarcolaenaceae respectively, in the Paris herbarium. For the (COETZEE & MÜLLER 1984), indicating that the “Material examined” cited below under each family was more widespread in the past, and that species, abbreviations are as follows: PN = Parc its current status as a Malagasy endemic is a result National; RNI = Réserve Naturelle Intégrale; RS of extinction elsewhere. = Réserve Spéciale; STF = Station Forestière. A Within Sarcolaenaceae, Eremolaena, Pentach- full listing of exsiccatae for each species, with laena and Perrierodendron appear to form a complete localities and latitude/longitude coordi- related and probably monophyletic group. As nates, has been compiled for the Madagascar indicated by CAPURON (1970), they share a num- Conspectus Project (SCHATZ et al. 1996), and is ber of diagnostic features, including free stipules available on the World Wide Web through W3 and inflorescence bracts, an involucre that is TROPICOS (http://mobot.mobot.org/Pick/Search/ reduced at anthesis (but which in certain species pick.html). Specimen data and images can also be expands somewhat in fruit), and seeds with little accessed through the SONNERAT database at or no endosperm and a smooth integument. (http://www.mnhn.fr/base/sonnerat.html). Moreover, members of all three genera share a Additional images are available on the Web at striking vegetative resemblance and are often dif- (http://mobot.mobot.org/MOBOT/Madagasc/ ficult to distinguish when sterile. Based on these sarco.html). Geographic coordinates indicated and other characters (especially of the pollen), in square brackets were assigned post facto CAPURON (1970) placed Eremolaena, Penta- using available information on Malagasy place chlaena and Perrierodendron in one of the infor- names and topographic maps, compiled as a mal groups he recognized within Sarcolaenaceae, gazetteer of botanical collecting localities in an interpretation supported by a preliminary Madagascar. ADANSONIA, sér. 3 • 2000 • 22 (1) 13 Lowry II P.P. et al. TAXONOMIC TREATMENT Key to the genera 1. Fruit indehiscent, subtended by an accrescent, entire involucre; ovary 2-carpellate ............ Perrierodendron 1’. Fruit dehiscent, the involucre accrescent or not, lobed; ovary 3-5-carpellate ................................................2 2. Sepals strongly unequal, the outer 2 much smaller; petals strongly contorted in bud; ovary 3-carpellate; ovules 2 per locule ................................................................................................................................ Eremolaena 2’. Sepals more or less equal in size; petals slightly contorted in bud; ovary 5-carpellate;
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