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.     

Isolation and characterization of nuclear microsatellite loci in the tropical arboreal palm Oenocarpus bacaba (Arecaceae)

Studies of mating patterns are needed to determine how habitat heterogeneity created by deforestation influences tropical plant populations. Eight microsatellite loci were isolated with a subtractive hybridization method for Oenocarpus bacaba (Arecaceae), a subcanopy palm tree at the Biological Dynamics of Forest Fragments Project, near Manaus, Brazil. Additionally, two heterospecific loci originally developed for the heart palm Euterpe edulis were shown to be variable. Loci averaged 9.6 alleles per locus. Five loci did not fit Hardy–Weinberg expectations with significant deficits of heterozygous genotypes consistent with null alleles. Many loci had high average probabilities of paternity exclusion.     

A systematic histological study of palm fruits. V. Subtribe Archontophoeniciae (Arecaceae)

Pericarp histology in the Archontophoenicinae provides little to characterize the subtribe as a whole, revealing instead two separate trends with parallels in other subtribes of the Areceae. The data support a close relationship among the three genera occurring in New Caledonia:Chambeyronia, Actinokentia, andKentiopsis, in which there is a complex endocarp consisting of short, oblique fibrous bundles embedded in a thick mantle of brachysclereids, and a loose endocarp of heavily fibrous, flattened vascular bundles adjacent to a relatively thin locular epidermis. The data also support a close relationship between the two genera of the New Zealand/Tasman Sea region:Hedyscepe andRhopalostylis, in which the pericarp is more or less fibrous throughout, with purely fibrous bundles in the outer pericarp and heavily fibrous vascular bundles in the inner pericarp. These results confirm relationships revealed by other morphological data.Archontophoenix appears to be most like the New Caledonian genera in its pericarp structure, with a similar mantle of short fibrous bundles embedded in a a mantle of brachysclereids in the outer pericarp, although it differs significantly in other aspects of morphology and anatomy.     

A systematic histological study of palm fruits. IV. Subtribe Oncospermatinae (Arecaceae)

Pericarp histology in the Oncospermatinae reveals distinctive characters at the generic level but no unique characters at the subtribal level. Pericarp structure is diverse but parallels the diversity found in other subtribes of the Areceae. The subtribe appears to be divisible into two distinct groups: one in which sclereids alone provide the primary protective barrier in the outer fruit and one in which fibrous bundles, some-times in combination with brachysclereids, form the outer barrier. This division of the subtribe supports similar conclusions derived from morphological data.     

A systematic histological study of palm fruits. III. Subtribe Iguanurinae (Arecaceae)

This study represents a preliminary sampling of the pericarp histology of the subtribe Iguanurinae (tribe Areceae, subfamily Arecoideae) of the family Arecaceae. At least one sample from each of the 27 recognized genera was examined and illustrated with a line drawing. This sampling serves to characterize fruit structure in the subtribe as a whole, to illustrate the diversity of pericarp adaptations found in the subtribe, to characterize the monotypic genera, to provide hypotheses about the characterization of the larger genera, and to test existing phylogenetic hypotheses about the Iguanurinae. There are no unique tissues present in the pericarp in this subtribe, but genera can be readily characterized by unique combinations and distributional patterns in common tissues. These patterns, and some prominent evolutionary trends, parallel those in related subtribes of Areceae, such as the Ptychospermatinae and Arecinae. Significant in this subtribe is variation in the distribution of tanniniferous cells, raphide-bearing cells and brachysclereids, in the sculpturing of the seed and the locular epidermis, in the thickness of the locular epidermis, in the thickness of the fibrous vascular bundle sheaths, and especially in the number, orientation and distribution of nonvascular fibrous bundles. One major trend is the formation of systems of separate fibrous bundles and their progressive displacement toward the outer layer of the fruit, where a complex exocarp may form. The diversity of pericarp structure in the Iguanurinae is far greater than in the two subtribes previously studied.     

A systematic histological study of palm fruits. VI. Subtribe Linospadicinae (Arecaceae)

The pericarp structure of representative species of the four genera of the subtribe Linospadicinae is described and compared. Tissues found in the pericarp of this group are similar to those found in other subtribes of the Areceae, with no characters unique to the subtribe and nothing to suggest its closest affinities. The four genera, as well as each species examined, do show distinctive pericarp features, however.Laccospadix andLinospadix are similar, both with a single series of fibrous, bundles and an outer series of prominent raphide-bearing cells dominating the pericarp. In bothHowea andCalyptrocalyx, a complex exocarp forms from a series of fibrous bundles and brachysclereids, but each genus has other distinctive characters.Howea has vascular bundles in the exocarp zone, an outer series of raphide-bearing cells, and a conspicuously thickened locular epidermis. Based on a limited sample of four species, raphide-bearing cells appear to be always interior to the exocarp inCalyptrocalyx, and the locular epidermis is thin.     

Functional and historical drivers of leaf shape evolution in palms (Arecaceae)

Aim

Leaves display a remarkable variety of shapes, each with potential ecological advantages in specific climates. While the relations between leaf shape and either climate or height have been relatively well studied in eudicots, the macroecological drivers of shape remain poorly known in monocots. Here, we investigated the associations between climate and plant height with the evolution of leaf shape in a clade with high species and morphological diversity.

Location

Global.

Time period

Cretaceous to contemporary.

Major taxa studied

Palms (Arecaceae).

Methods

We apply a Bayesian phylogenetic mixed model to test for associations between climate and leaf shape (all – entire-leaved, pinnate-dissected, palmate-dissected and costapalmate). We further reconstruct the ancestral leaf shape using multistate speciation and extinction models and compare the frequency of shapes with global temperatures through time.

Results

We find that plant height associates with dissected leaves and that annual precipitation associates with pinnate shapes. The ancestral leaf shape is unclear, but early diversification was dominated by pinnate-dissected palms, which has remained the most species-rich form of leaves throughout palm history.

Main Conclusions

Palms that are tall and live in humid regions are more likely to have pinnate leaves. Through geological time scales, temperature did not play an obvious role in determining leaf shapes. This study contributes to our understanding of how the diversity of leaf shapes is linked to biological and climatic factors.     

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.     

Comparative anatomy and systematics of Senghas's cushion species of Maxillaria (Orchidaceae)

Using data obtained through anatomy and morphology, we used cladistics to examine the monophyly of Senghas's proposed classification of Maxillaria cushion plants and his placement of Mormolyca ringens. Trignidium obtusum was chosen as the outgroup. Leaves have multicellular hairs sunken in crypts, primarily anomocytic or primarily tetracytic stomatal apparatuses, homogeneous mesophyll, and scattered fibre bundles. Three types of adaxial hypodermis were observed: (1) water-storage cells, (2) fibre bundles scattered among water-storage cells, and (3) fibre bundles scattered among chlorenchymatous cells. Abaxial hypodermis of fibre bundles occurs in several Maxillaria species and in Trigonidium obtusum. At the midvein of the leaf, adaxial mesophyll cells of most species are anticlinally extended and empty, and the abaxial mesophyll is usually collenchymatous. Vascular bundles are collateral and usually in a single series. Pseudobulb epidermal cell walls are thin, or outer walls are thickened. Ground tissue consists of water-storage and assimilatory cells with vascular bundles and associated lacunae scattered throughout. Roots are velamentous and exodermal cell walls are usually n-thickened with tenuous bands of scalarifom thickenings on longitudinal walls. Tilosomes may be plaited, baculate, or spongy. Endodermal cell walls are usually U-thickened and pericycle cell walls are usually O-thickened opposite phloem sectors. Stegmata line the periphery of the thickened pericycle cells opposite phloem sectors in M. picta. Pith may be parenchymatous or sclerenchymatous. According to our phylogenetic analysis, Mormolyca ringens is consistently nested within the cladistic structure of Maxillaria. Therefore, Maxillaria likely is paraphyletic if Mormolyca ringens is recognized as generically distinct. It appears that Senghas's subgroup divisions of the unifoliate pseudobulbous maxillarias may also be artificial.     

Leaf morphology and anatomy of Camellia section Camellia (Theaceae)

The delimitations of species in Camellia section Camellia have been disputed for many years, resulting from uncertain relationships among species. Leaf morphological and anatomical characters for 54 species and three varieties in this section were investigated to reveal the relationships. Principal component analysis and cluster analysis were conducted using the transformed data for quantitative and qualitative characters from leaf morphology and anatomy. Combining the results of statistical analysis with comparative leaf characters of morphology and anatomy, we discussed the taxonomic treatment of section Camellia by Chang compared with that of Ming and we conclude that section Camellia consists of c. 50 species. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 456–476.     

Isolation of 23 polymorphic microsatellite loci in the Neotropical palm Oenocarpus bataua Martius (Arecaceae)

We report the isolation of 23 microsatellite loci obtained from a (GA)_n‐enriched genomic library of Oenocarpus bataua var. bataua. The average number of alleles per locus, the mean observed and expected heterozygosities for these microsatellite loci revealed a high level of variability. The transferability of the developed markers to other Oenocarpus species and other genera within the Euterpeae tribe was high, except for the Hyospathe genus.     

The shoot apical meristem of oil palm (Elaeis guineensis; Arecaceae): developmental progression and dynamics

Background and Aims

Oil palm, an unbranched perennial monocotyledon, possesses a single shoot apical meristem (SAM), which is responsible for the initiation of the entire above-ground structure of the plant. To compare the palm SAM structure with those of other monocots and to study variations in its structure throughout the life of the plant, its organization was characterized from the embryonic stage to that of the reproductive plant.

Methods

SAM structure was studied by a combination of stained histological sections, light and confocal microscopy, and serial section-based three-dimensional reconstructions.

Key Results

The oil palm SAM is characterized by two developmental phases: a juvenile phase with a single tunica-corpus structure displaying a gradual increase in size; and a mature phase characterized by a stable size, a modified shape and an established histological zonation pattern. In mature plants, fluctuations in SAM shape and volume occur, mainly as a consequence of changes in the central zone, possibly in relation to leaf initiation.

Conclusions

Development of the oil palm SAM is characterized by a juvenile to mature phase transition accompanied by establishment of a zonal pattern and modified shape. SAM zonation is dynamic during the plastochron period and displays distinct features compared with other monocots.     

Evolution of reproductive traits in the mahagony family (Meliaceae)

Meliaceae are a mostly pantropical family in the Sapindales, bearing flowers typically provided with a staminal tube, formed by filaments that are fused partially or totally. Nevertheless, several genera of subfamily Cedreloideae have free stamens, which may be adnate to an androgynophore in some taxa. The fact that the family exhibits a wide diversity of floral and fruit features, as well as of sexual systems and pollination syndromes, presents interesting questions on the evolutionary processes that might have taken place during its history. In this study, we analyzed the distribution of 20 reproductive morphological traits of Meliaceae, upon an available molecular phylogenetic framework, using 31 terminals from the family's two main clades (Cedreloideae and Melioideae), plus six Simaroubaceae taxa as outgroup. We aimed to identify and/or confirm synapomorphies for clades within the family and to develop hypotheses on floral evolution and sexual systems in the group. Our reconstruction suggests that the ancestor of Meliaceae was possibly provided with united stamens and unisexual flowers in dioecious individuals, with a subsequent change to free stamens and monoecy in the ancestor of Cedreloideae. Most characters studied show some degree of homoplasy, but some are unique synapomorphies of clades, such as the haplostemonous androecium. An androgynophore defines the Cedrela‐Toona clade. The comparative approach of our study and the evolutionary hypotheses generated herein reveal several aspects demanding further structural investigation, and possible evolutionary pathways of the reproductive structures along with the lineages' diversification, mostly related to the specialization of sexual systems, floral biology, and dispersal strategies.     

The occurrence and structure of apparently bisexual flowers in the date palm, Phoenix dactylifera L. (Arecaceae)

Individuals of Phoenix dactylifera L. have expanded pistillodes or pseudocarpels in staminate flowers. These pseudocarpels are located in the centre of the male flowers and are surrounded by stamens. The gynoecium has the characteristic three carpellate arrangement commonly found in female date palm flowers. Pseudocarpels from male flower buds can expand into parthenocarpic fruit. Histology of the expanded pistillodes or pseudotarpels is similar to that of normal carpels from pistillate plants. These pseudocarpels lack ovules. Nutrient medium containing 10 mg 1^-1 of 2,4-dichlorophenoxyacetic acid or p-chlorophenylacetic acid and 0.3% activated neutralized charcoal enhanced the development and outgrowth of the pseudocarpels of cultured male flowers.     

Evaluation of the bromatological potential of seeds and fruits ofSabal Mexicana Mart. (Arecaceae)

This work presents bromatological analyses performed on seeds and fruits of Sabal mexicana Mart.; a comparison with data existent of other palms and some phytochemical data are presented. A discussion of the possible importance of some results obtained and the potential use of this palm is presented

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Evaluacion del potencial bromatologico de semillas y frutos de sabal mexicana, mart. (arecaceae)

Résumé  Este trabajo presenta análisis bromatolólogicos realizados en semillas y frutos de Salbal mexicana Mart. Los resultados se comparan con datos existentes para otras palmas y se presentan algunos datos fitoquimicos; ademús se discute acerca de la posible importancia de algunos de los resultados obtenidos y su uso potential

     

Genetic structure among morphotypes of the endangered Brazilian palm Euterpe edulis Mart (Arecaceae)

Euterpe edulis (Arecaceae) Mart has high ecological and economic importance providing food resources for more than 58 species of birds and 20 species of mammals, including humans. E. edulis is the second most exploited nontimber product from Brazilian Atlantic Forest. Due to overexploitation and destruction of habitats, E. edulis is threatened by extinction. Euterpe edulis populations have large morphological variations, with individuals having green, red, or yellow leaf sheath. However, no study has related phenotypic distinctions between populations and their levels of genetic structure. Thus, this study aimed to evaluate the diversity and genetic structure of different E. edulis morphotypes. We sampled 250 adult individuals in eight populations with the different morphotypes. Using 14 microsatellite markers, we access genetic diversity through population genetic parameters calculated in the GenAlex program and the diveRsity package in R. We used the Wilcoxon test to verify population bottlenecks and the genetic distance of Nei and Bayesian analysis for genetic clusters. The eight populations showed low allele richness, low observed heterozygosity, and high inbreeding values (f). In addition, six of the eight populations experienced genetic bottlenecks, which would partly explain the low genetic diversity in populations. Cluster analysis identified two clusters (K = 2), with green morphotype genetically distinguishing from yellow and red morphotypes. Thus, we show, for the first time, a strong genetic structure among E. edulis morphotypes even for geographically close populations.     

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.     

Comparative floral structure and systematics in the Asian palm genus Rhapis (Arecaceae,Coryphoideae)

A comparative study of the floral structure in the species of the genus Rhapis (Arecaceae, Coryphoideae, Rhapidinae) is presented. Flowers are mainly unisexual, with three sepals, three petals, 6 stamens or staminodes and three carpels or carpellodes. Some evidences of basal congenital and apical postgenital fusion of the carpels, first time reported in the genus, were observed in the gynoecium. Ovules are basally attached and crassinucellate; they appear to be slightly anatropous. The morphology of the filaments suggests a division of the species into two groups: Rhapis excelsa and R. subtilis exhibit thick and keeled filaments, whereas R. gracilis, R. humilis, R. laosensis, R. micrantha and R. multifida have slender, non-keeled filaments. Relationships of Rhapis with the rest of the genera of Rhapidinae are inferred on the light of floral structure.