Comparative morphology of female flowers and systematics in Geonomeae (Arecaceae)

Female floral structure is compared in Geonomeae (Arecaceae). A perianth is formed by two alternate whorls of three basally congenitally united and imbricate sepals and three basally congenitally united and apically valvate petals. A sterile androecium is formed by a variable number of staminodes, which are united into a tube. The gynoecium shows three more or less equally developed carpels or is pseudomonomerous (Geonoma). The single anatropous ovule per carpel is median, either basal or at mid-height of the ovary. A septal nectary is present at the base and mid-height of the ovaries and exits at different levels of the ovary. Carpels in pseudomonomerous gynoecia seem to be basistylous, but the styles are more lateral or apical in gynoecia with all three carpels equally developed. Stigmas expose unicellular or multicellular (Welfia) papillae at anthesis. Pollen tube transmitting tracts and a compitum are present in the ventral slits of the postgenitally united styles. Floral structure in Geonomeae is compared with other Arecaceae, especially Arecoideae, in a morphological and systematic context.     

Phylogenetic relationships among arecoid palms (Arecaceae: Arecoideae)

Background and Aims

The Arecoideae is the largest and most diverse of the five subfamilies of palms (Arecaceae/Palmae), containing >50 % of the species in the family. Despite its importance, phylogenetic relationships among Arecoideae are poorly understood. Here the most densely sampled phylogenetic analysis of Arecoideae available to date is presented. The results are used to test the current classification of the subfamily and to identify priority areas for future research.

Methods

DNA sequence data for the low-copy nuclear genes PRK and RPB2 were collected from 190 palm species, covering 103 (96 %) genera of Arecoideae. The data were analysed using the parsimony ratchet, maximum likelihood, and both likelihood and parsimony bootstrapping.

Key Results and Conclusions

Despite the recovery of paralogues and pseudogenes in a small number of taxa, PRK and RPB2 were both highly informative, producing well-resolved phylogenetic trees with many nodes well supported by bootstrap analyses. Simultaneous analyses of the combined data sets provided additional resolution and support. Two areas of incongruence between PRK and RPB2 were strongly supported by the bootstrap relating to the placement of tribes Chamaedoreeae, Iriarteeae and Reinhardtieae; the causes of this incongruence remain uncertain. The current classification within Arecoideae was strongly supported by the present data. Of the 14 tribes and 14 sub-tribes in the classification, only five sub-tribes from tribe Areceae (Basseliniinae, Linospadicinae, Oncospermatinae, Rhopalostylidinae and Verschaffeltiinae) failed to receive support. Three major higher level clades were strongly supported: (1) the RRC clade (Roystoneeae, Reinhardtieae and Cocoseae), (2) the POS clade (Podococceae, Oranieae and Sclerospermeae) and (3) the core arecoid clade (Areceae, Euterpeae, Geonomateae, Leopoldinieae, Manicarieae and Pelagodoxeae). However, new data sources are required to elucidate ambiguities that remain in phylogenetic relationships among and within the major groups of Arecoideae, as well as within the Areceae, the largest tribe in the palm family.     

A revision of the Greater Antillean species of Bactris (Bactridinae: Arecaceae)

Three species ofBactris are recognized in the Greater Antilles:B. cubensis (from Cuba),B. plumeriana (from Hispaniola), andB. jamaicana (from Jamaica). A cladistic analysis of the non-ocreate clade—i.e.,Bactris cubensis, B. plumeriana, B. jamaicana, B. macana, andB. gasipaes—confirmed that the Greater Antillean species ofBactris form a monophyletic group, i.e., the Antillean clade. Synapomorphies supporting the Antillean clade are mesocarp and endocarp fibers numerous, parallel, predominantly narrow, few broad, extending the entire length of the endocarp; and leaf segments 45–80 per side of rachis. The presence of fiber-sclereids in the leaf lamina, and petals of the staminate flowers ovate to ovatetrullate, with sparsely branched fibers, may represent additional synapomorphies.Bactris plumeriana andB. jamaicana are hypothesized to be sister species, a grouping weakly supported by their short anthers. Each species has at least one autapomorphy; therefore, all are considered to be a cladospecies. Noteworthy interpopulational variation occurs withinBactris plumeriana andB. cubensis.     

A systematic histological analysis of palm fruits VII. The Cyrtostachydinae (Arecaceae)

Fruit specimens representing five taxa of the genusCyrtostachys were examined histologically in order to characterize the pericarp anatomy of the monogeneric subtribe Cyrtostachydinae (tribe Areceae, subfamily Arecoideae), as part of an ongoing survey of the family. The pericarp in this genus can be characterized by a combination of papillate epidermis, heavy layer of tanniniferous/pigmented cells below the epidermis, a system of vascular bundles with thick fibrous sheaths with purely fibrous bundles frequently above and below, absence of brachysclereids, and a very thin sclerified locular epidermis. On the basis of pericarp structure alone, the genus might be most closely related to theGronophyllum alliance of the subtribe Arecinae. This diverges somewhat from the hypothesis of relationship with theAreca group of the Arecinae resulting from two DNA-based phylogenetic studies, and even further from the hypothesis of relationship withIguanura suggested by another DNA-based phylogenetic study.     

A revision of the genus Sclerosperma (Arecaceae)

Summary  A taxonomic revision of the palm genus Sclerosperma (Arecaceae) is presented. Three species are recognised: S. mannii H. Wendl., which is relatively widespread from Liberia to the Democratic Republic of Congo; S. walkeri A. Chev., which is apparently confined to the interior of Gabon and a band along the Congo River; and S. profiziana, a new species previously considered conspecific to S. mannii that is found in southwest Ghana, Congo, the Democratic Republic of Congo and Angola. The taxonomic history, morphology, distribution and conservation status of the genus and each species are discussed.     

Early inflorescence and floral development in Cocos nucifera L. (Arecaceae: Arecoideae)

Palms are generally characterized by a large structure with a massive crown that creates difficulties in anatomical studies. The flowering behaviour of palm species may be a useful indicator of phylogenetic relationships and therefore evolutionary events. This paper presents a detailed histological study of reproductive development in coconut (Cocos nucifera L.), from initiation up to maturation of staminate and pistillate flowers. Reproductive development in coconut consists of a sequence of individual events that span more than two years. Floral morphogenesis is the longest event, taking about one year, while sex determination is a rapid process that occurs within one month. The inflorescence consists of different ultimate floral structural components. Pistillate flowers are borne in floral triads that are flanked by two functional staminate flowers. The staminate flowers are born in floral diads towards the base of the rachilla followed by solitary flowers in the middle to top of the rachilla. Three primary phases were identified in reproductive development, namely, transition of axillary bud into inflorescence bud, formation of floral buds, and sexualisation of individual flower buds. All developmental events with respect to stage or time of occurrence were determined.     

A new species of Bactris (Palmae) from French Guiana

A new species,Bactris nancibaensis, from French Guiana, is described and illustrated.     

Floral structure and development in the monoecious palm Gaussia attenuata (Arecaceae; Arecoideae)

Background and Aims

Sexual dimorphism, at both the flower and plant level, is widespread in the palm family (Arecaceae), in contrast to the situation in angiosperms as a whole. The tribe Chamaedoreeae is of special interest for studies of the evolution of sexual expression since dioecy appears to have evolved independently twice in this group from a monoecious ancestor. In order to understand the underlying evolutionary pathways, it is important to obtain detailed information on flower structure and development in each of the main clades.

Methods

Dissection and light and scanning electron microscopy were performed on developing flowers of Gaussia attenuata, a neotropical species belonging to one of the three monoecious genera of the tribe.

Key Results

Like species of the other monoecious genera of the Chamaedoreeae (namely Hyophorbe and Synechanthus), G. attenuata produces a bisexual flower cluster known as an acervulus, consisting of a row of male flowers with a basal female flower. Whereas the sterile androecium of female flowers terminated its development at an early stage of floral ontogeny, the pistillode of male flowers was large in size but with no recognizable ovule, developing for a longer period of time. Conspicuous nectary differentiation in the pistillode suggested a possible role in pollinator attraction.

Conclusions

Gaussia attenuata displays a number of floral characters that are likely to be ancestral to the tribe, notably the acervulus flower cluster, which is conserved in the other monoecious genera and also (albeit in a unisexual male form) in the dioecious genera (Wendlandiella and a few species of Chamaedorea). Comparison with earlier data from other genera suggests that large nectariferous pistillodes and early arrest in staminode development might also be regarded as ancestral characters in this tribe.     

A phylogenetic analysis of the Euterpeinae (Palmae; Arecoideae; Areceae) based on morphology and anatomy

The Euterpeinae contains six neotropical genera. There has been continual disagreement on generic and subgeneric boundaries in the subtribe.Euterpe andPrestoea, andJessenia andOenocarpus, have been repeatedly united and separated. A phylogenetic analysis based on 54 morphological and anatomical characters gave one tree of 127 steps.Euterpe is separate fromPrestoea, butJessenia andOenocarpus are best treated as one genus. Subgeneric relationships ofEuterpe andOenocarpus are also analyzed and discussed.     

New species of <Emphasis Type="Italic">Geonoma</Emphasis> (Palmae) from Ecuador

Five new species of Geonoma from Ecuador, G. awaensis, G. ecuadoriensis, G. hollinensis, G. lanata, and G. skovii, are described and illustrated and are compared to similar species. Their distributions are mapped.     

A phylogenetic analysis of the palm subtribe Oncospermatinae (Arecaceae) based on morphological characters

Subtribe Oncospermatinae (Arecaceae: Arecoideae: Areceae) is a diverse group of spiny Old World palms. The subtribe includesOncosperma, a widespread Asian genus of five species, along with seven monotypic genera, all endemic to the Seychelles and Mascarene Islands of the western Indian Ocean. A phylogenetic analysis was conducted in order to test the monophyly of subtribe Oncospermatinae with respect to other Old World genera of tribe Areceae. A matrix of 38 morphological characters was scored for 29 taxa, including 11 species of the Oncospermatinae. A single most parsimonious tree was found, resolving the subtribe as a polyphyletic group of two distinct clades. One clade containingAcanthophoenix, Deckenia, Oncosperma, andTectiphiala was placed as sister to a large group that includes members of subtribes Archontophoenicinae, Arecinae, Iguanurinae, and Ptychospermatinae. The other clade of Oncospermatinae, including the Seychelles endemic generaNephrosperma, Phoenicophorium, Roscheria, andVerschaffeltia, was resolved as sister to the Madagascar endemic subtribe Masoalinae, and may have arisen in the western Indian Ocean region.     

Phylogenetic utility of the nuclear gene malate synthase in the palm family (Arecaceae)

There is currently a shortage of DNA regions known to be useful for phylogenetic research in palms (Arecaceae). We report the development and use of primers for amplifying and sequencing regions of the nuclear gene malate synthase. In palms the gene appears to be single-copy, with exon regions that are phylogenetically informative within the family. We constructed a phylogeny of 45 palms and five outgroup taxa using 428 bp of malate synthase exon regions. We found that some major clades within the family were recovered, but there was a lack of resolution among the genera in subfamilies Arecoideae, Ceroxyloideae, Coryphoideae, and Phytelephantoideae. In a second analysis, malate synthase exon regions totaling 1002 bp were sequenced for 16 palms and two outgroup taxa. There was increased bootstrap support for some groups and for the placement of the monotypic genus Nypa as sister to the rest of the family. A comparison with data sets from noncoding regions of the chloroplast genome indicates that malate synthase sequences are more variable and potentially contain more phylogenetic information. We found no evidence of multiple copies of the malate synthase gene in palm genomes.     

Calospatha subsumed in Calamus (Arecaceae: Calamoideae)

Summary  Based on previously published phylogenetic research, the genus Calospatha Becc. (Calamoideae) is placed in synonymy within Calamus L. The new combination, Calamus calospathus (Ridl.) W. J. Baker & J. Dransf. is made.     

Pathways of karyological differentiation in palms (Arecaceae)

Karyological data are given for 56 palm taxa coming from all 6 palm subfamilies. In 11 genera and 17 species, chromosome numbers are reported for the first time. Most chromosome numbers in palms range between 2n = 36 and 2n = 26 in dysploid series. Species of the same genus usually exhibit identical chromosome numbers which additionally may be constant in larger groups of closely related genera (Coryphoideae trib.Corypheae with nearly always 2n = 36,Arecoideae subtribesEuterpeinae andRoystoneinae with 2n = 36,Arecoideae subtrib.Butiinae with mostly 2n = 32). Polyploidy among palms is of minor significance but the endemic Madagascan genusVoanioala (2n = 606 ± 3) is the most striking exception. — With respect to structure of interphase nuclei and longitudinal differentiation of prophase and metaphase chromosomes, the palm family is highly differentiated. Euchromatin types with different prophase condensation properties and fluorochrome and C-banding patterns of heterochromatin permit a discrimination of several subfamilies on the nuclear level (Arecoideae, Ceroxyloideae, Nypoideae, Phytelephantoideae, Calamoideae).Arecoideae andCeroxyloideae, andNypoideae andPhytelephantoideae have some features in common. Subfam.Coryphoideae s. l. is a non-uniform group. — Nuclear characters among palms exclusively found in recentCoryphoideae subtrib.Thrinacinae link palms with other monocotyledons. Most probably, such a nuclear condition represents an ancestral state in the evolution of palm genomes within subfam.Coryphoideae s. l., but also the conspicuous nuclear characters of the other modern palm subfamilies appear to be derived from a similar starting point, since transitional character states are still present in subfam.Calamoideae and some taxa of subfam.Arecoideae. Early karyoevolution in palms obviously did not involve numerical change of the ancient chromosome number of 2n = 36 which started subsequently, as a dysploid reduction in numerous parallel series, independent in subfam.Coryphoideae (2n = 36 to 2n = 28),Calamoideae (2n = 36 to 2n = 26),Ceroxyloideae (2n = 34 to 2n = 26), andArecoideae (2n = 36 to 2n = 28). Possible mechanisms of karyological change are discussed. — Karyological characters are compared to morphological, ecological, taxonomical, and chorological features, and give some new insight into older and more recent phases of palm evolution. (1) Strong deviations in vegetative or floral morphology are often accompanied by major karyological differences, and sometimes the direction of advancement can be traced through intermediate stages. (2) Apart fromCoryphoideae subtrib.Thrinacinae, the strongest concentration of apparently original karyological traits is found in the more basal members of each subfamily. (3) The most successful and actively radiating colonizers of the forest floors in evergreen tropical forests which belong to completely different subfamilies (Old WorldLicuala, New WorldChamaedorea andGeonoma), appear to be very advanced karyologically.     

Classification of specimens in the Geonoma stricta (Palmae) complex: the problem of leaf size and shape

Specimens representingGeonoma stricta, a widespread species in the Amazon region, are variable morphologically, especially in leaf size and shape. Ordination and clustering of quantitative morphological variables indicate that the specimens can be classified in at least four different ways, but none of these is fully supported by Discriminant Analysis. The problem revolves around a group of specimens with intermediate leaf size and shape. Mapping of specimens shows that no grouping is geographically separate from any other. An infraspecific classification is not possible based on the data analyzed here. Principal Component Analysis and a test of hybridity indicate that a small percentage of specimens might be of hybrid origin.