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 (: 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 and highlight unexpected affinities between the tribe Leopoldinieae and representatives of the western Pacific Ocean tribe Pelagodoxeae.

The , 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 and supported by the supertree of Baker et al. (2009). However, the The number of 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. /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, and (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 (Putz 1979), whereas genera Pelagodoxa and , 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 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 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. 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. material studied Species Collection Reproductive status Repository Leopoldinia piassaba Guanchez s.n . (19 Oct. 1990) Fruits L.H. Bailey Wallace Guanchez s.n . (22 Oct. 1990) Fruits Hortorium Guanchez s.n . Pre-anthetic ♀ buds Guanchez 5080 Late ♀ buds Guanchez 5081 ♀ buds

Leopoldinia pulchra Guanchez 4912 Early fruits L.H. Bailey Mart. Guanchez 4911 Early fruits Hortorium Guanchez s.n . (06 Oct. 1990) ♂ buds Guanchez s.n . Early ♂ and ♀ buds

Guevara 79a ♂ buds Venezuelan Guevara 79b Early ♂ and ♀ buds Central Univ. Guevara 78 Early ♂ buds

Manicaria saccifera Stauffer (Feb. 2012) Late ♀ and ♂ buds Conservatory Gaertn. and Botanical Garden of Geneva

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1. Habit of Leopoldinia pulchra . 2. Habit of L. piassaba . 3. Stem surface with leaf sheaths of L. pulchra . 4. Infructescence of L. pulchra . 5. of L. pulchra . 6. Fences constructed with stems of L. pulchra by indigenous tribes of the Orinoco region. (photos: L. Guevara and F. Stauffer) at the Natural History Museum of Geneva, female flowers are solitary or arranged in triads Switzerand. with two male flowers (Fig. 14) flanking one Results central female . Solitary female flowers are usually more robust than the ones borne Flower morphology and anatomy of in combination with male flowers. When not Leopoldinia in floral triads, male flowers are either solitary The inflorescence is interfoliar and bears or inserted in dyads; they are surrounded by a unisexual flowers sunken in shallow pits. The brown, dense tomentum (Fig. 13).

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The male flowers are sessile, oblong, ovoid, 1 idioblasts were observed in the mesophyll of mm long or even smaller in diameter, in all the gynoecium, especially from the base up to cases smaller than the pistillate flowers. The the mid-height of the ovary. Tanniferous calyx reaches one-third of the flower length idioblasts were also observed in the pistillode and is composed of three distinct, imbricate, of the male flowers. Raphide idioblasts were keeled and striate sepals, with toothed margins. not seen in the tissues studied. The corolla is composed of three distinct, valvate, triangular-ovate petals, with entire Flower morphology and anatomy of margins. The petals are on the inner side Manicaria marked by the impression of the anthers. The The inflorescence is interfoliar and bears androecium is composed of six, latrorse, unisexual flowers sunken in pits. The dorsifixed stamens (Figs. 15 & 16). Three staminate flowers are solitary and spirally stamens are antepetalous and three arranged on the entire rachillae, whereas only antesepalous. The ones in the antesepalous few pistillate flowers concentrate at the base of position are attached at a slightly lower level the rachilla. than the antepetalous. The filaments are short, broad and connate only at the very base; they The male flowers are oblong, asymmetrical, 5 have inflexed tips. The anthers are short and mm long and 4 mm wide. The calyx is ovate in a cross-section (Fig. 17). The pistillode composed of three slightly imbricate, distinct has a three-lobed barrel shape. sepals, which are membranous, irregular in size and shape, slightly keeled, with fibrous The female flowers are sessile, globose, 2.5–3 variously notched margins. The corolla is mm long and 2.5–3 mm in diameter at late composed of three, very thick valvate petals bud stage. The calyx is composed of three (Fig. 25), one remarkably smaller than the distinct, imbricate, rounded and membranous others. The petals are distinct and adnate to sepals, with slightly toothed margins. The the base of the filaments of the peripheral corolla is composed of three distinct, valvate, stamens. The androecium is polyandrous (Figs. rounded petals, with entire margins. The sterile 21, 22, 24), with ca. 20–25 introrsely dehiscent, androecium is represented by six reduced dorsifixed stamens; the length of the filaments staminodes, which are short, flat and truncate. and the anthers, including the shape of the The gynoecium is synascidiate, composed of latter, vary according to the position of the three postgenitally united carpels, which at stamen in the androecium. Two filaments mid-height of the ovary become separated. No connate up to the level of the anthers were epithelium that could be associated with a observed in some flowers; pistillode lacking. septal nectary was identified in the carpel flanks. The gynoecium is trilocular and The pistillate flowers are oblong, ovoid, 9 mm triovulate; however, only one ovule fully long and 6 mm in diameter, slightly develops towards anthesis. The stage of compressed and asymmetrical. The calyx is development of the ovule characterizes the composed of three imbricate sepals, which are shape of the gynoecium, ranging from slightly distinct, membranous and irregular in size and pyramidal to rounded (Figs. 18, 19). The ovule shape, with fibrous margins. The corolla is is campylotropous, bitegmic, crassinucellate composed of three very thick petals, two and hanging; it is laterally attached to the notably larger than the remaining one. The ventral side of the locule and fills the entire petals are valvate with an obtuse apex and cavity (Fig. 20). Towards the micropyle the congenitally fused up to 4–5 mm, with entire outer and inner integuments are 8 and 4 cell margins. Staminodes 9, small, thin, basally layers thick, respectively; the micropyle is adnate to the ovary, upwards free; each straight and slightly expanded due to the large staminode is served by one central vascular obturator. The three stigmatic branches are bundle; six staminodes are antepetalous and lined with unicellular papillae. The pollen tube oblong in cross-section, whereas three are transmitting tract (PTTT) separates downwards antesepalous and triangular in cross-section in three branches, corresponding to the three (Fig. 28). The gynoecium is obovoid, composed locules. It reaches the ovule by surrounding the of three congenitally united carpels; all of funiculus. them are equally developed at late bud stage, giving to the gynoecium a triangular shape in The base of the female flower is characterized cross-section (Fig. 29). The ovary region is by a six cell layers thick transition zone of ovoid, topped with a pyramidal stigma with square or rhomboid cells that may be three clearly differentiated branches; the style interpreted as an abscission zone. Tanniferous is short. The PTTT is common at the stigmatic-

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7. Habit of Manicaria saccifera . 8. Leaves; 9. Infructescence. 10. Detail of a cross-sectioned . 11. Portion of inflorescence with male flowers at late bud. 12. Thatching constructed by the Warao Indians in the Orinoco Delta. (photos: L. Guevara and F. Stauffer) style region and separates downwards in three the region of the chalaza (Fig. 26). Tannins branches, one directing each locule. It reaches idioblasts concentrate at the base of the the ovule by surrounding the funiculus. The gynoecium but were also observed throughout gynoecium is triloculate and triovulate. The the gynoecium, around the three ovules. These ovule is anatropous, bitegmic and tanniferous idioblasts were not observed in the crassinucellate (Fig. 23). It is attached to the ovary walls. The lack of tannins in the ovary ventral side and in the upper half of the locule walls enables to distinguish two types of and fills the entire cavity. mesophylls in the cross-section of the gynoecium (Fig. 27). Raphide idioblasts were The whole gynoecium is highly vascularized, not seen in the tissues studied. especially in the peripheral zones cor- responding with the thick ovary walls (Fig. Discussion 27). Each carpel is served by one dorsal This paper represents a contribution to the vascular bundle, 10–12 lateral procambrial knowledge of the floral structure in the tribes strands and four ventral vascular bundles. Manicarieae and Leopoldinieae. The floral Vascular bundles were also clearly observed in structure of Leopoldinia and Manicaria had

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13 –20. Morphology and anatomy of the flowers in Leopoldinia . 13. Portion of rachilla showing developing male floral buds, note dense hairs covering the rachilla, scale bar = 250 μm; 14. Portion of rachilla showing a triad composed of one central female flower and two male flowers at bud stage, scale bar = 250 μm; 15. Side view of a male flower at late bud stage, perianth removed, scale bar = 125 μm; 16. Longitudinal section of a male flower at late bud stage, perianth removed, scale bar = 100 μm; 17. Cross-section of an anther, note the well differentiated endothecium, scale bar = 50 μm; 18. Upper view of female flower at late bud stage, petals removed, scale bar = 500 μm; 19. Longitudinal section of a female flower at early bud stage, scale bar = 500 μm; 20. Longitudinal section of an ovule, note micropylar region facing the ventral side of the locule, scale bar = 100 μm. never previously been studied in detail, in spite composed of one central female flower and of the large economic importance of these two lateral male flowers sunken in pits. This palms for indigenous populations in the type of floral arrangement unequivocally neotropics and the fact that both taxa display supports the inclusion of these two tribes unclear phylogenetic relationships among the within the subfamily Arecoideae. The two taxa of the core arecoid clade. genera display an important number of shared Leopoldinia and Manicaria are characterized by floral characters, namely oblong staminate floral triads, at least at the base of the rachillae, flowers, with three distinct and imbricate

187 PALM S Villímová & Stauffer: Floral Structure Vol. 57(4) 2013

21 –29. Morphology and anatomy of the flowers in Manicaria . 21. Side view of a polyandrous male flower, perianth removed, scale bar = 500 μm; 22. Upper view of a polyandrous male flower, perianth removed, scale bar = 500 μm; 23. Detail of an ovule, arrow pointing to the micropyle region, scale bar = 100 μm; 24. Cross- section of a male flower showing anthers, scale bar = 250 μm; 25. Cross- section of an anther with a detail on the mesophyll of the petal, scale bar = 100 μm; 26. Cross-section of an ovule showing vascular bundles emerging from the funiculus, scale bar = 100 μm; 27. Cross-section of the ovary, at the left side mesophyll with tanniferous cells, at the right side thick ovary wall with large vascular bundles, scale bar = 100 μm; 28. Cross-section of the basal part of the ovary showing two staminodes with different shapes, triangular in antesepalous position and oblong in antepetalous position, scale bar = 250 μm; 29. Cross-section of the ovary showing three equally developed ovules, scale bar = 500 μm. sepals with variously notched or toothed gynoecium presents distinct staminodes and a margins and three valvate petals. The pistillate syncarpous, triloculate, triovulate ovary topped flowers are characterized by three distinct and by three stigmatic branches. Contrary to what imbricate sepals and three valvate petals; the has been observed in other arecoid groups

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(Rudall et al. 2003, Stauffer et al. 2003, Stauffer stamens in Manicaria is characteristic of the et al. 2004) septal nectaries were not detected male flowers, and female flowers display an in either of the two genera, suggesting that increased number of staminodes. Leopoldinia other possible types of reward attract potential has consistently six stamens in the male pollinators (e.g. pollen). On the other hand, flowers and six staminodes in the female raphide-containing idioblasts, suggested to be flowers. a reward for bee-pollinators (D’Arcy et al. 1996) or as defense devices (Sakai et al. 1972), were Our study explored the possible taxonomic not evident in any organs of the flowers. The relationships among representatives of the five presence of tanniferous idioblasts throughout tribes of the basal core arecoids, based on the gynoecium in both genera suggests that comparison with data presented in similar these inclusions may be regarded as a defense studies of floral structure (Table 2). Indeed, mechanism against herbivores. Interestingly, representatives of all five tribes share only a the absence of the septal nectaries and raphide- few common characters, namely distinct containing idioblasts in Leopoldinia and imbricate sepals, valvate petals, inflexed Manicaria appears to be unique, compared with stamens at bud stage in the male flowers and representatives of other basal core arecoids imbricate sepals and three differentiated (Table 2). stigmatic branches in the female flowers. Morphological and anatomical characters In spite of several shared characters, the flowers associated with the floral structure of of Leopoldinia differ from Manicaria in several Leopoldinia show surprising affinities with important aspects concerning the floral flowers of the palm genus Pelagodoxa (Table structure. Male flowers of Leopoldinia are 2), endemic to the Marquesas Islands, and characterized by the typical supports the relationships proposed by the trimery, represented by six latrorse, basally molecular studies of Norup et al. (2006) and connate stamens with broad filaments, and a Baker et al. (2011). Leopoldinia and Pelagodoxa three-lobed syncarpous pistillode. In the case present oblong male flowers, distinct and of Manicaria, the male flowers have 20–25 thin, imbricate sepals, triangular-ovate valvate elongated and introrse stamens, and the petals, six dorsifixed stamens and a syncarpous pistillode is completely lacking. The stamen pistillode. Female flowers of these two genera filaments are rarely connate up to the anthers have distinct and imbricate sepals, distinct and the filaments of the peripheral stamens are petals and more or less rounded, basally connate to the petals. The female pseudomonomerous, triloculate, triovulate flowers of the two genera differ mostly in the gynoecia, topped by three stigmas covered shapes of the gynoecium and perianth parts. with unicellular papillae. Leopoldinia is characterized by a pyramidal to rounded gynoecium, rounded petals and Leopoldinia resembles Euterpe only with respect sepals. Manicaria has on the contrary an to perianth characters. These two genera have obovoid gynoecium with oblong or irregular similar distinct, rounded, variously tattered or and asymmetrical sepals and petals. notched, imbricate sepals, distinct, valvate petals, six wide stamens inflexed at bud stage Although both genera have syncarpous, and three-lobed pistillode in male flowers. trilocular and triovulate gynoecia, the Female flowers of Leopoldinia and Euterpe have gynoecium of Manicaria presents three equally similar, distinct, imbricate sepals and distinct developed ovaries at late bud stage, whereas petals. Flowers of Leopoldinia share with the gynoecium of Leopoldinia shows what may the presence of distinct and imbricate be described as late pseudomonomery, in sepals, valvate petals and three-lobed pistillode which the fertile ovary and the two sterile in male flowers and imbricate sepals, valvate ovaries are almost equal in size and shape (Stauffer & Endress 2003). The ovules are petals, pseudomonomerous triloculate campylotropous and hanging in Leopoldinia , gynoecium with papillate stigmatic branches whereas they are anatropous and laterally and six staminodes in the female flowers. attached in Manicaria . In the case of Manicarieae does not show evident affinities Leopoldinia , the globose gynoecium is topped with any other representative of the basal core by three free stigmatic branches with arecoids. The surprisingly isolated position of unicellular papillae, whereas in Manicaria, the this palm, already suggested by the molecular obovoid gynoecium is topped by a rather study of Baker et al. (2011), may be supported smooth pyramidal stigmatic region with by the male flower with many stamens but connate stigmatic branches. Numerous lacking a pistillode and a clearly trimerous

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191 PALM S Villímová & Stauffer: Floral Structure Vol. 57(4) 2013 gynoecium in the female flower. The presence BAKER , W.J., M.V. N ORUP , J.J. C LARKSON , T.L.P. of a completely enveloping prophyll and COUVREUR , J.L. D OWE , C.E. L EWIS , J.-C. peduncular bract (even at fruiting stage) may PINTAUD , V. S AVOLAINEN , T. W ILMOT AND M.W. be added as a unique character for Manicaria CHASE . 2011. Phylogenetic relationships among the basal core arecoid tribes. Male among arecoid palms (Arecaceae: flowers of Manicaria and Pelagodoxa have an Arecoideae). Annals of Botany 108: oblong shape, distinct and imbricate sepals, 1417–1432. distinct petals and dorsifixed introrse stamens. The female flowers of these two taxa share BAKER , W.J., V. S AVOLAINEN , C.B. A SMUSSEN -L ANGE , distinct, imbricate sepals, a triloculate, M.W. C HASE , J. D RANSFIELD , F. F OREST , M.M. triovulate gynoecium with anatropous ovules. HARLEY , N.W. U HL AND M. W ILKINSON . 2009. Manicaria shares with Euterpe the same group Complete generic-level phylogenetic of characters that is shared with Leopoldinia , analyses of palms (Arecaceae) with namely distinct, imbricate sepals with comparisons of supertree and supermatrix variously notched margins, distinct, valvate approaches. Systematic Biology 58: 240–56. and asymmetrical petals in male flowers, and BERNAL , R. AND G. G ALEANO . 2010. Notes on distinct, imbricate sepals and an ovoid Mauritiella , Manicaria and Leopoldinia . Palms gynoecium in the female flowers. The male 54: 119–132. flowers of Manicaria and Geonoma share distinct sepals, valvate petals and introrse DRANSFIELD , J., N.W. U HL , C.B. A SMUSSEN , W.J. stamens, whereas the female flowers of both BAKER , M.M. H ARLEY AND C.E. L EWIS . 2008. genera have imbricate sepals, valvate, basally Genera Palmarum: the Evolution and united petals and triloculate gynoecium with Classification of Palms. Kew Publishing, an anatropous ovule. Finally, flowers in London. representatives of Geonomateae and Euterpeae show some affinities with Manicarieae and D’A RCY , W.G., R.C. K EATING AND L.S. B UCHMANN . Leopoldinieae, but further studies may be 1996. The calcium oxalate package or so- necessary to infer more defined relationships called resorption tissue in some angiosperm between these groups. anthers. Pp 159–191 in D’A RCY , W.G. AND R.C. K EATING (eds.) The Anthers: Form, Acknowledgments Function and Phylogeny. Cambridge We thank the L.H. Bailey Hortorium (Cornell University Press, New York. University, Ithaca) and Lorena Guevara GAERTNER , J. 1791. De Fructibus et Seminibus (Venezuelan Central University) for providing Plantarum. Stuttgart. valuable material for the present study. We express special acknowledgements to André HAHN , W.J. 2002. A phylogenetic analysis of Piuz (Natural History Museum of Geneva) for the Arecoid Line of palms based on plastid his guidance and help with SEM images and DNA sequence data. Molecular Phylogenetics to Michèle Crèvecoeur (University of Geneva) and Evolution 23: 189–204. for her assistance with the micrographs. HENDERSON , A. 1995. The Palms of The Amazon. Literature Cited Oxford University Press, Inc. ASMUSSEN , C.B. AND M.W. C HASE . 2001. Coding HENDERSON , A. 2011. A revision of Leopoldinia and noncoding plastid DNA in palm (Arecaceae). Phytotaxa 32: 1–17. systematics. American journal of botany 88: 1103–1117. IGERSHEIM , A. AND O. C ICHOCKI . 1996. A simple ASMUSSEN , C.B., J. D RANSFIELD , V. D EICKMANN , method for microtome sectioning of A.S. B ARFOD , J.-C. P INTAUD AND W.J. B AKER . prehistoric charcoal specimens, embedded 2006. A new subfamily classification of the in 2-hydroxyethyl methacrylate (HEMA). palm family (Arecaceae): evidence from Review of Palaeobotany and Palynology 92: plastid DNA phylogeny. Botanical Journal 389–393. of the Linnean Society 151: 15–38. KUCHMEISTER , H., I. S ILBERBAUER -G OTTSBERGER AND BAKER , W.J. AND T.L.P. C OUVREUR . 2012. Global G. G OTTSBERGER . 1997. Flowering, pollination, biogeography and diversification of palms nectar standing crop, and nectaries of Euterpe sheds light on the evolution of tropical precatoria (Arecaceae), an Amazonian rain lineages. I. Historical biogeography. Journal forest palm. Plant Systematics and Evolution of Biogeography 40: 274–285. 206: 71–97.

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