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.     

Multiplicity in palm uses by the Huaorani of Amazonian Ecuador

An ethnobotanical study of the palms used by the Huaorani in the Yasuní National Park and Huaorani Ethnic Reserve in Amazonian Ecuador was carried out. In this inventory, 37 palm species were found; all were used by the Huaoranis. One hundred and ninety-one different uses were recorded in eight ethnobotanical categories. Most species (64.9%) were used for house construction and human food. More than half of the species were used for domestic utensils (59.4%) and hunting and fishing implements (54%). A comparison is made between these data and past studies for the other six indigenous communities from Amazonian Ecuador. This paper shows the highest diversity of useful palm species and the highest number of different uses ever recorded for an indigenous group in Amazonian Ecuador. The data combine quantitative and qualitative approaches.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 149–159.     

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.     

Traditional morphometrics in plant systematics and its role in palm systematics

A review is given of the role of traditional morphometrics in plant systematics. The three most commonly used techniques of data analysis – Cluster Analysis, Principal Component Analysis and Discriminant Analysis – are discussed. The kinds of data that can be taken from palm specimens and the problems of using specimens as data sources are outlined. Published systematic studies of palms using traditional morphometrics are reviewed. More recent studies indicate that: hybrid zones between species may be common; infraspecific diversity is greater than previously suspected; there may be more than double the currently accepted number of species; and our current knowledge of morphological variation in palms is superficial. A procedure for scientific systematics is given, which incorporates traditional morphometric methods.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 103–111.     

Pollination in Geonoma macrostachys and Three Congeners,G. acaulis,G. gracilis,and G. interrupta

A continuous 15 month study of the floral ecology of four syntopic understorey palm species of Genoma was conducted in Amazonian Peru lowland rainforest. The spicate inflorescences of G. macrostachys, G. acaulis and G. gracilis are strictly protandrous and the plants are functionally dioecious. Data suggest that in G. macrostachys and G. acaulis pollination is based on a mimicry system, the pistillate flowers mimicking the staminate ones in colour, shape and scent. Pollen-collecting meliponine bees (Hymenoptera, Apidae, Meliponinae) and pollen-feeding syrphid flies (Diptera, Syrphidae) which visit inflorescences during both sexual stages are the pollinators of G. macrostachys. Geonoma acaulis is pollinated by small pollen-feeding weevils (Coleoptera, Curculionidae, Derelomini) that visit male and female spikes. Additionally, in G. macrostachys another pollinator type, viz. euglossine bees (Hymenoptera, Apidae, Euglossinae), which are attracted and rewarded by both types of flowers may account for long-distance pollination. The palm G. gracilis shows a very distinct pollination system. Although opportunistic insect visitors are attracted to the inflorescences of this species it seems to be mainly anemophilous because pollen becomes powdery during an thesis. The branched inflorescences of G. interrupta are also protandrous, but unlike the other species of Geonoma observed, staminate and pistillate anthesis of individual flowers are, for the most, overlapping. A broad spectrum of visitors is attracted (bees, wasps, flies, and beetles), which all may act as pollinators. Outcrossing is especially encouraged during the purely female phase at the end of the flowering cycle when there are no more staminate flowers in the inflorescence. Effects on the reproductive biology and population structure of different pollination systems and breeding system are discussed.     

Fossil palm flowers in Dominican and Mexican amber

Twelve palm flowers in one piece of Dominican amber and a single flower in a second piece are described as Trithrinax dominicana sp. nov. (Thrinacinae: Coryphoideae), representing the first fossil record of this genus. There are no members of Trithrinax in the Greater Antilles today. Two staminate flowers in Mexican amber are described as Socratea brownii sp. nov. (Iriarteinae: Arecoideae) and represent the first fossils of this genus. A third palm flower in Mexican amber, possibly belonging to the subtribe Euterpeinae, is characterized. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 57–61.     

Cellular ultrastructure and crystal development in Amorphophallus (Araceae)

Background and Aims: Species of Araceae accumulate calcium oxalate in the form ofcharacteristically grooved needle-shaped raphide crystals andmulti-crystal druses. This study focuses on the distributionand development of raphides and druses during leaf growth inten species of Amorphophallus (Araceae) in order to determinethe crystal macropatterns and the underlying ultrastructuralfeatures associated with formation of the unusual raphide groove. Methods: Transmission electron microscopy (TEM), scanning electron microscopy(SEM) and both bright-field and polarized-light microscopy wereused to study a range of developmental stages. Key Results: Raphide crystals are initiated very early in plant development.They are consistently present in most species and have a fairlyuniform distribution within mature tissues. Individual raphidesmay be formed by calcium oxalate deposition within individualcrystal chambers in the vacuole of an idioblast. Druse crystalsform later in the true leaves, and are absent from some species.Distribution of druses within leaves is more variable. Drusesinitially develop at leaf tips and then increase basipetallyas the leaf ages. Druse development may also be initiated incrystal chambers. Conclusions: The unusual grooved raphides in Amorphophallus species probablyresult from an unusual crystal chamber morphology. There aremultiple systems of transport and biomineralization of calciuminto the vacuole of the idioblast. Differences between raphideand druse idioblasts indicate different levels of cellular regulation.The relatively early development of raphides provides a defensivefunction in soft, growing tissues, and restricts build-up ofdangerously high levels of calcium in tissues that lack theability to adequately regulate calcium. The later developmentof druses could be primarily for calcium sequestration.     

Fossil palm flowers in Dominican and Baltic amber

Five palm flowers in Dominican amber and one in Baltic amber are described or characterized. Palaeoraphe dominicana gen. et sp. nov. in the subtribe Livistoninae, is described from one perfect flower in Dominican amber. Roystonea palaea sp. nov. is described from one staminate and one pistillate flower in Dominican amber. Three other palm flowers, two perfect flowers from Dominican amber and one staminate flower from Baltic amber, are briefly characterized and figured.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 361–367.     

Crystal types and their distribution in the bark of African genistoid legumes (Fabaceae tribes Sophoreae,Podalyrieae, Crotalarieae and Genisteae)

The occurrence and distribution of seven crystal types in 114 bark samples from 25 genera and 91 species, representing all four tribes of African genistoid legumes, are reported. The seven types are prismatic, druse (including irregular crystal clusters), navicular (including truncated navicular, here reported for the first time), spherical crystal cluster, styloid, crystal sand and acicular crystals in sheaf‐like aggregates. Unlike most studies, the elemental composition of the crystals was examined using X‐ray microanalyses. With the exception of acicular crystals, all crystals showed the typical peaks of calcium (sometimes accompanied by small amounts of magnesium). Acicular crystals in sheaf‐like aggregates were composed only of carbon and oxygen, indicating that they are organic and precipitate during drying or after fixation in alcohol. These crystals are found only in the two early‐diverging lineages of Podalyrieae (Cyclopia and Virgilia+Calpurnia clades), consistent with the phylogenetic pattern in the tribe (indicating a secondary loss). Navicular crystals are restricted to Podalyrieae and Crotalarieae. Prismatic crystals in bark are proposed to be the ancestral condition, with multiple losses (or reversals) in Podalyrieae, Crotalarieae and Genisteae. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 178 , 620–632.     

Sporogenesis and gametogenesis in Sladenia and their systematic implication

Sporogenesis and gametogenesis in the monotypic genus Sladenia is reported for the first time. The anther wall is of the monocotyledonous type of formation and consists of four-layers, namely a glandular tapetum, one middle layer, an endothecium, and an epidermis. Fibrous thickenings are developed in the epidermis at or near the stage of anther maturity rather than in the endothecium. Successive cytokinesis in microsporogenesis results in tetrahedral or rarely isobilateral tetrads of microspores. Pollen grains are two-celled when shed. The ovule is bitegmic, tenuinucellate, and anatropous. The nucellus degenerates during meiosis of the megaspore mother cell. The micropyle is formed by only the inner integument. Development of the female gametophyte conforms to the Adoxa type. Sladenia has been treated either as a member of Actinidiaceae, Dilleniaceae or Theaceae s.l. , or as belonging to an independent family, Sladeniaceae. Recent molecular studies have suggested that it belongs to Ternstroemiaceae within the Ericales s.l . Several noteworthy embryological characters are distinct from those of the other putatively related families and support the recognition of a separate family, Sladeniaceae. Additional embryological evidence confirms this conclusion.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 305–314.     

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.     

Water balance in the arborescent palm, Sabal palmetto. I. Stem structure, tissue water release properties and leaf epidermal conductance

The functional importance of water storage in the arborescent palm, Sabal palmetto, was investigated by observing aboveground water content, pressure-volume curve parameters of leaf and stem tissue and leaf epidermal conductance rates. The ratio of the amount of water stored within the stem to the leaf area (kg m^?2) increased linearly with plant height. Pressure-volume curves for leaf and stem parenchyma differed markedly; leaves lost turgor at 0.90 relative water content and –3.81 MPa, while the turgor loss point for stem parenchyma occurred at 0–64 relative water content and ?0.96 MPa. Specific capacitance (change in relative water content per change in tissue water potential) of stem parenchyma tissue was 84 times higher than that of leaves, while the bulk modulus of elasticity was 346 times lower. Leaf epidermal conductance rates were extremely low (0.32–0.56 mmol m^?2 s^?1) suggesting that S. palmetto are able to strongly restrict foliar water loss rates. Structurally, stems of S. palmetto appear to be well suited to act as a water storage reservoir, and coupled with the ability to restrict water loss from leaf surfaces, may play an important role in tree survival during periods of low water availability.     

Osmotic responses of preimplantation mouse and bovine embryos and their cryobiological implications

Cells subjected to the events occurring before, during, and after freezing and thawing are exposed to major changes in the osmotic pressure of the surrounding medium; i.e., the osmolalities can exceed 30. An important question in understanding the mechanisms of injury is whether cells respond as ideal osmometers to these strongly anisotonic solutions. Mouse and bovine embryos from eight-cell to blastocyst stage were used to investigate the question. They were found to behave as ideal osmometers at room temperature over a wide range of tonicities; i.e., from four times isotonic to almost 1/3 times isotonic, ideality being defined by a Boyle-van't Hoff equation. Embryo volumes increased from 40 to 200% of isotonic over this range and survivals of mouse embryos were unaffected. However, outside this range the membrane apparently becomes leaky and the survival of mouse embryos drops sharply. Osmolalities rise to high values during freezing and the paper develops the thermodynamic equations to show how computed cell volumes as a function of subzero temperature can be translated into the Boyle-van't Hoff format of cell volume as a function of the reciprocal of osmolality.     

Ontogenetic development in architecture and biomass allocation of 13 rattan species in Indonesia

Rattans are climbing, nonbranching palms with diverse growth forms ranging from stems that remain at the forest floor to stems that reach the canopy. We analyzed changes in architecture and biomass allocation during ontogenetic development of 13 Indonesian species in the genera Calamus, Ceratolobus, Daemonorops, Korthalsia, and Plectocomiopsis. Species included both nonclimbers (<5 m tall maximum) and climbers (10 to 50 m tall). Nonclimbers retain a rosette form—that is, thick, short internodes with length/internode diameter ratio <10 but with long petioles—throughout their lives. Leaf and internode shapes of climbers at the early stage resemble those of nonclimbers, but internodes later become longer (internode length/diameter ratio ≥10) and petioles become shorter and developed one of two kinds of climbing organs at the adult stage. These developmental changes reduce self-shading within the crown. Some climbers have dwarf blades at the early stage and skip the rosette form. Principal component analysis of biomass allocation indicated that growth strategies to attain the adult stage are diverse. These results suggest that rattans reach maturity at different phases along a series of ontogenetic development stages and generate diverse growth forms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Palm snorkelling: leaf bases as aeration structures in the mangrove palm (Nypa fruticans)

Mangrove species have evolved specialized structures, such as pneumatophores, to supply oxygen to the roots, but, in Nypa fruticans, the only mangrove palm, no such structure has been reported. This study aimed to determine the adaptations of N. fruticans to the mangal habitat with special reference to the air‐supplying structure. Following senescence, the rachis is abscised at the zone of junction with the leaf base. Simultaneously, lenticels develop so that, when abscission is completed, a network of mature lenticels covers the leaf base. Expansigenous aerenchyma with increasing porosity towards the stem junction occurs in the leaf base. The first two root branching orders present a subero‐lignified rhizodermis and exodermis, and the cortex consists of schizo‐lysigenous aerenchyma with wide lacuna, limiting radial oxygen loss and facilitating longitudinal oxygen transport to living tissues. Lifespan estimation suggests that leaf bases can live for up to 4 years following abscission, ensuring the persistence of aeration structures. This study provides structural evidence indicating that N. fruticans has evolved a unique type of air‐supplying structure in the mangal habitat. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 257–270.     

Development of the gametophytes, flower, and floral vasculature in Cochliostema odoratissimum (Commelinaceae)

Flowers of Cochliostema odoratissimum are trimerous with three fertile stamens, three unequal antherless staminodes. and three connate carpels. The fertile stamens are on one side of die flower and united by their filaments, forming a compound structure that curves to the flower's right as the flower opens. The thecae arc longitudinally dehiscent, spirally coiled, and enveloped by pctaloid extensions of the filaments of the two lateral stamens contributing to the three-staminate structure. Anther wall development is of the monocotyledonous type. Tapetal raphides are formed and appear to be widespread in CCommelinaceae. Also known from Philydraceae and. perhaps. Haemodoraccae, tapetal raphides and their taxonomic distribution may be of phylogenetic utilitv. Microsporogcnesis is successive, forming both isobilateral and decussate tetrads. Pollen is shed as single binucleatc grains. Each ovary locule contains ten to twelve hemianatropous, crassinucellar. bitegmic ovules on axile placentae. The micropyle is formed by both the inner integument and one side of the outer integument. Megagamctophyte development is of the Polygonum type. The mature megagamctophyte consists of an egg apparatus, fusion nucleus, and three antipodals. the latter showing signs of degeneration. The salient features of the floral vasculature are the same as in the few other commclinaceous species for winch complete data are available. Relative to the floral vasculature in the other species, differences in the vasculature lie primarily in the presence and origin of lateral carpel bundles and in the number of sepal and ovule traces.     

Changes in the distribution of melanophores and xanthophores inTriturus alpestris embryos during their transition from the uniform to banded pattern

Summary The change in distribution of melanophores from stage 28+ (uniform melanophore pattern) to stage 34 (banded melanophore pattern) and the participation of xanthophores in these changes has been investigated inTriturus alpestris embryos by studying the social behaviour of single cells. While melanophores are clearly visible from outside the embryo at stage 28+, xanthophores cannot be recognized from the outside until after stage 34. In ultrathin sections of stage 34 embryos, xanthophores are observed alternating with melanophores in a zonal distribution (Epperlein 1982). Using detached pieces of dorsolateral trunk skin, which retain their chromatophores after detachment, the entire distribution of melanophores and xanthophores can be visualized in a scanning electron microscope (SEM). In ambiguous cases (early stages), cells were reprocessed for transmission electron microscopy (TEM) and the presence of the characteristic pigment organelles was assessed. In addition, xanthophores were specifically identified by pteridine fluorescence with dilute ammonia. Pteridines were also identified chromatographically in skin homogenates. The combination of these methods allowed precise identification and quantitative determination of melanophores and xanthophores. Both cell types were present as codistributed, biochemically differentiated cells at stage 28+. Changes in the pattern up to stage 34 were due to the rearrangement at the epidermal-mesodermal interface of a relatively fixed number of melanophores which became preferentially localised at the dorsal somite edge and at the lateral plate mesoderm, and to the distribution of an increasing number of xanthophores to subepidermal locations in the dorsal fin and between the melanophore bands. Concomitant was an increase in the thickness of the epidermal basement membrane and a change in shape of chromatophores from elongate via stellate to rosette shaped, which may be correlated with a shift from migratory to sessile phases.     

Changes in lipid profiles in the developing haustorium of oil palm seeds

Germinated oil palm seeds were placed in special trays in the field and at different intervals the haustoria were harvested. Changes in haustorial lipids were followed until the eighth week after germination. The moisture content decreased while the lipids increased. The haustorial fatty acid profile was similar to that of the palm kernel. Changes in fatty acid composition at different weeks after germination were observed. The major haustorial lipids were triacylglycerols; free fatty acids and diacylglycerols were also present     

New costapalmate palm leaves from the Oligocene Ningming Formation of Guangxi,China, and their biogeographic and palaeoclimatic implications

New palm leaves from the Oligocene Ningming Formation are placed into the morphogenus Sabalites because of their costapalmate leaf shape. Four taxa are described on the basis of leaf compressions with cuticular structure. S. guanxiensis sp. nov. is characterised by hypostomatic leaf blades with a stout costa and a symmetrical base. Sabalites cf. asymmetricus has amphistomatic leaf blades with a long, delicate costa and an asymmetrical base. Sabalites sp. 1 has amphistomatic leaf blades with a long, massive costa and wide segments. Sabalites sp. 2 is characterized by hypostomatic leaf blades with a prominent costa and an asymmetrical base. The four new palm taxa expand our understanding of the floristic elements and features of the Oligocene Ningming flora. Together with the other three palm taxa that were previously reported from the Ningming Formation, our material indicates that the Oligocene Ningming flora had a rich diversity of costapalmate palms. The relatively high species diversity of palms and other plants corroborate that the Oligocene Ningming flora represents a warm and humid climate.