PALM S Villímová & Stauffer: Floral Structure Vol. 57(4) 2013 VERONIKA VILLÍMOVÁ Ecole Normale Supérieure de Lyon, Floral Structure 15 parvis René Descartes – BP 7000 69342 Lyon in the Cedex 07, France veronika.villimova@gmail. Neotropical com Tribes AND FRED W. S TAUFFER Conservatoire et Jardin Leopoldinieae botaniques de la Ville de Genève, and Manicarieae Université de Genève, Laboratoire de (Arecaceae: systématique végétale et biodiversité, ch. de l’Impératrice 1, Arecoideae) case postale 60, 1292 Chambésy/Genève, Switzerland Floral structure is studied in detail for the first time in the monogeneric neotropical tribes Leopoldinieae and Manicarieae (Arecaceae). In order to infer taxonomic relationships of these groups, morphological and anatomical data in members of the two tribes are compared with available studies in other tribes of the basal clades of the core arecoids, as defined by recent molecular phylogenies. Our results suggest an isolated position for Manicaria and highlight unexpected affinities between the tribe Leopoldinieae and representatives of the western Pacific Ocean tribe Pelagodoxeae. The genus Leopoldinia , dedicated to the of the tribe Manicarieae, as defined by archduchess of Austria, Josefa Carolina Dransfield et al. (2008) (Figs. 7–12). These two Leopoldina, was described in 1824 by the remarkable monogeneric tribes belong to the celebrated palm botanist Carl F. P. von Martius large subfamily Arecoideae, known to be the (1794–1868) (Figs. 1–6). It represents the single largest and most diverse subfamily within representative of the neotropical tribe palms (Dransfield et al. 2008). The tribes Leopoldinieae (Martius 1824). The genus Manicarieae and Leopoldineae represent two Manicaria was described in 1791 by the basal lineages of the core arecoid clade as German botanist Joseph Gaertner (1732–1791) defined by Baker and Couvreur (2012), here and corresponds with the single representative called “basal core arecoids,” including also PALMS 57(4): 181 –193 181 PALM S Villímová & Stauffer: Floral Structure Vol. 57(4) 2013 tribes Pelagodoxeae, Euterpeae and Geo- suggested also by Asmussen et al. (2006) in nomateae. the Plastid DNA tree and supported by the supertree of Baker et al. (2009). However, the The number of species within Leopoldinia and prk -based phylogeny of Lewis and Doyle (2002) Manicaria remains a subject of taxonomic and the combined prk & rpb2 tree of Baker et debate. In the case of Leopoldinia, different al. (2011) placed the Manicarieae as sister to a numbers of species have been proposed since clade composed of the tribes Euterpeae, the description of the genus. Hence, Martius Leopoldinieae, Pelagodoxeae, Geonomateae (1824) recognized two species, whereas Wallace and included the tribe Leopoldinieae in their (1853) increased the number to three, a plastid DNA tree for the first time in a taxonomic point of view that was supported molecular phylogeny of the palm family. In by Henderson (1995). Bernal and Galeano this analysis the tribe was resolved as sister to (2010) recognized only two species and more the Manicarieae. Later, the Leopoldinieae was recently, Henderson (2011) proposed that all resolved as sister to Euterpeae (Lewis & Doyle species published so far for the genus (L. 2002, Loo et al. 2006) and as sister to the piassaba Wallace, L. major Wallace and L. Areceae/Euterpeae, Geonomateae and pulchra Mart.) should be recognized as valid. Manicarieae clades in the plastid DNA tree of A similar situation can be traced in Manicaria, Hahn (2002). A sister relationship of a genus described by Gaertner (1791) based on Leopoldinieae with the Manicarieae/ only one species ( Manicaria saccifera Gaertn). Geonomateae clade was indicated by Different taxonomic and floristic treatments Asmussen et al. (2006) and recovered by the on the genus have recognized three species study of Baker et al. (2009). In all these (i.e. Wessels Boer 1988) or only one (i.e. phylogenies the relationships of the tribe Henderson 1995). More recently, Bernal and Leopoldinieae with other arecoid groups was Galeano (2010) have recognized M. martiana always resolved with low bootstrap support. Burret and M. saccifera Gaertn. as the only two Norup et al. (2006) proposed a sister accepted species for the genus. relationship, although with moderate Both genera are distributed in the neotropical bootstrap support, between Leopoldinieae and region. Manicaria has a wide distribution, Pelagodoxeae, the latter endemic to the ranging from Central America, across Trinidad, Marquesas Islands and the western half of the the Orinoco Delta and the Guianas to the mainland New Guinea. The same relationship lower Amazon River (Dransfield et al. 2008). was recovered by Baker et al. (2011), with In contrast, Leopoldinia is an endemic palm strong bootstrap support. This relationship is genus of the Amazon basin, restricted to the surprising because both tribes, Leopoldinieae Rio Negro and upper Orinoco region of and Pelagodoxeae, display highly disjunct and Venezuela, Colombia and Brazil (Dransfield et geographically isolated distribution patterns al. 2008, Henderson 1995, Stauffer 2000). (Dransfield et al. 2008). The most recent Representatives of both genera have been molecular phylogenetic analysis of the palm reported to be economically important for family confirms previous studies in which indigenous groups of the Amazon and Orinoco Leopoldinieae is resolved as sister to a clade basins. At least two species of Leopoldinia (L. composed of the tribes Geonomateae and piassaba , L. pulchra ) have been reported as Manicarieae (Baker & Couvreur 2012). The Old economically important for their stems (Fig. 6), World tribe Pelagodoxeae, composed of the fibers, leaves and fruits (Putz 1979), whereas genera Pelagodoxa and Sommieria, remains the leaves of Manicaria are used primarily for relatively close in the topology presented. thatching (Dransfield et al. 2008) by ethnic Morphology and anatomy of palm groups such as the Warao Indians from the reproductive structures have been studied in Orinoco delta region (Fig. 12). detail in only 4% of the almost 2500 palm The molecular phylogenetic relationships of species (Stauffer et al. 2002). Floral structure, Leopoldinieae and Manicarieae with the especially anatomy has never been studied in remaining tribes of Arecoideae are not yet detail in Leopoldinia and Manicaria . Infor- completely understood. Both tribes are mation on gross morphological characters has included in the “core” arecoid clade, but their been provided for members in all tribes of the precise position remains unclear (Dransfield basal core arecoids (Dransfield et al. 2008), but et al. 2008). The Manicarieae was found to be the floral structure has been studied in detail sister to the Geonomateae in the RFLP tree of in only three of them: Pelagodoxeae (Stauffer Uhl et al. (1995); this relationship was et al. 2004), Geonomateae (Stauffer & Endress 182 PALM S Villímová & Stauffer: Floral Structure Vol. 57(4) 2013 2003, Stauffer et al. 2003), and Euterpeae collected in the Venezuelan Amazon. Male and (Kuchmeister et al. 1997). As a contribution female flowers at bud stage of M. saccifera were to the understanding of the reproductive collected by FWS in the Orinoco Delta structures in arecoid palms and in order to (Venezuela). explore relationships of the two enigmatic For the anatomical investigations, small monogeneric tribes Leopoldinieae and fragments of rachillae or individual flowers Manicarieae within the core arecoid clade, a were evacuated, dehydrated and embedded in thorough study of the floral structures of the resin Kulzer’s Technovit 7100 (2- Leopoldinia piassaba , Leopoldinia pulchra and hydroxyethyl methacrylate [HEMA]). Further Manicaria saccifera has been conducted. The details of this techniques are given in specific aims of the present study are (1) to Igersheim and Cichocki (1996). The material contribute to a better understanding of the was serially cross sectioned and longitudinally floral structure of the neotropical tribes sectioned at 7–10 μm using a rotary microtome Leopoldinieae and Manicarieae and (2) to (Leitz 1512), stained with ruthenium red and explore the systematic relationships of toluidine blue and mounted in Assistant- Leopoldinieae and Manicarieae with other Histokitt mounting medium. Observations and groups of the basal clade of the core arecoids photographs were made with a digital light using floral structural characters. microscope (NIKON Eclipse 80i) at the Materials and Methods Laboratory of Cytology and Vegetal Histology (University of Geneva); the permanent slides The morphological and anatomical study was were deposited at the Laboratory of Micro- based on flowers collected from wild Morphology of the Conservatory and populations and fixed in alcohol (Table 1). Botanical Garden of Geneva. Inflorescences at several stages of development and young infructescences of L. piassaba and For scanning electron microscopy (SEM), L. pulchra were collected by Dr. Lorena Guevara fragments of inflorescences, individual flowers (Venezuelan Central University) and also and individual floral organs were dehydrated, obtained from the spirit collection of the L. H. critical-point dried and sputter-coated with Bailey Hortorium (Cornell University, Ithaca, gold. Micrographs were obtained using a Zeiss USA). In both cases the material was originally DSM 940A SEM (Orion 6.60 Imaging System) Table 1. Plant material studied Species Collection Reproductive status Repository Leopoldinia piassaba Guanchez