The impact of utilization of palm products on the population structure of the vegetable ivory palm (Hyphaene petersiana,arecaceae) in North-Central Namibia

Indigenous trees fulfil many subsistence and economic needs in north-central Namibia.Hyphaene petersiana provides a range of products which contribute to most aspects of people’s livelihoods. Of particular importance is its income-generating capacity through the use of palm leaves for basket production and the sale of liquor distilled from the fruits. This study investigates the population structure ofHyphaene petersiana in two areas of different human and livestock densities. Data were recorded for height class distribution, basal diameter of mature, stemmed individuals and sex ratios. These parameters of population structure indicate a reduction in the recruitment of mature palms and an increase in single-stemmed, vegetatively reproduced palm suckers of the smallest size class (0.5 m). This trend is more pronounced in the site with greater human and livestock population densities. It appears to be related to high recorded levels of browsing by livestock of juvenile, unstemmed palms, despite the unpalatability of palm leaves. This acts to prevent recruitment into larger size classes and increase the compensatory growth of palm suckers, the latter being enhanced due to reduced competition through the prior removal by grazing animals of grasses and other herbaceous species. Accompanying this heavy pressure on juvenile palms are destructive uses of mature, stemmed palms, including their felling for construction purposes and tapping for palm wine. With regional human population increase, exacerbated by a recent trend to privatise land and raise pressure on remaining communal resources, it is possible that these destructive uses of mature palms will increase to unsustainable levels. Concern is thus expressed in this study regarding the long-term viability ofHyphaene petersiana populatiops in this area.     

Elephant herbivory of knob-thorn (Senegalia nigrescens) and ivory palm (Hyphaene petersiana) in Bwabwata National Park,Caprivi, Namibia: The role of ivory palm as a biotic refuge

African elephants have major impacts on vegetation, particularly at high densities. Knob-thorns (Senegalia nigrescens) are typically ring-barked by elephant, and high levels of mortality are common at high elephant densities. Our study aimed to test whether ivory palm clusters (Hyphaene petersiana) form a biotic refuge for knob-thorn against elephant herbivory. We measured the density, damage and mortality of knob-thorns in sites differing according to ivory palm presence and elephant density, and thus, the probability of knob-thorn encounter by elephants. The site with palms and low elephant density, had a high density of knob-thorns, but lower proportions of damaged and dead trees, than sites without palms but with similar or higher elephant density. In the former, knob-thorns were associated with palm clusters, particularly saplings and young adults. In this site, low proportions of damaged and dead knob-thorns were recorded in palm clusters, compared with outside clusters, and to those in the other sites. Our study also showed that juvenile palms which protected knob-thorns, suffered low mortality in contrast to subadult palms. We have no evidence but implicate elephants and suggest that in palm clusters, subadult palms are more accessible to elephants than knob-thorns because of the different methods of utilisation.     

Evolution of lamina anatomy in the palm family (Arecaceae)

The unique properties of tree building in Arecaceae strongly constrain their architectural lability. Potentially compensating for this limitation, the extensive diversification of leaf anatomical structure within palms involves many characters whose alternate states may confer disparate mechanical or physiological capabilities. In the context of a recent global palm phylogeny, we analyzed the evolution of 10 such lamina anatomical characters and leaf morphology of 161 genera, conducting parsimony and maximum likelihood ancestral state reconstructions, as well as tests of correlated evolution. Lamina morphology evolves independently from anatomy. Although many characters do optimize as synapomorphic for major clades, anatomical evolution is highly homoplasious. Nevertheless, it is not random: analyses indicate the recurrent evolution of different cohorts of correlated character states. Notable are two surface layer (epidermis and hypodermis) types: (1) a parallel-laminated type of rectangular epidermal cells with sinuous anticlinal walls, with fibers present in the hypodermis and (2) a cross-laminated type of hexagonal cells in both layers. Correlated with the cross-laminated type is a remarkable decrease in the volume fraction of fibers, accompanied by changes in the architecture and sheath cell type of the transverse veins. We discuss these and other major patterns of anatomical evolution in relation to their biomechanical and ecophysiological significance.     

Chloroplast DNA evolves slowly in the palm family (Arecaceae)

A survey of cpDNA restriction-site variation for 22 species representing five of the six subfamilies of the palm family was conducted. Phylogenetic reconstructions based on the restriction-site data are in general agreement with conventional analyses based on morphological characters. Base-substitution estimates, derived from the restriction-site data, indicate a potential 5-13-fold decrease in substitution rates within the palms relative to rate estimates for annual plant taxa. Approximately 1,000 bp of the rbcL gene (ribulose-1,5-bisphosphate carboxylase, large subunit) were sequenced from species representing two subfamilies of the palms. The complete DNA sequence data are in accord with the restriction-site data and indicate a total rate of nucleotide substitution that is about eightfold lower than that observed for annual plants.     

Wounaan and Emberá use and management of the fiber palmAstrocaryum standleyanum (Arecaceae) for basketry in eastern Panamá

This study examines the use of Astrocaryum standleyanum for basketry by Wounaan and Emberá artisans in Panama. Fibers from the palm are a traditional source of material for baskets, which have become commercial art items within the last fifteen years. Data on palm use were collected over two years, largely from Darién Province. Results indicate that the palm is destructively harvested, usually for a single spear leaf. Artisans remove the fibers from the pinnae and color them with dye plants. Astrocaryum standleyanum strands are sewn around a foundation of Carludovica palmata to make coiled baskets. Wounaan weavers are considered the better artisans, perhaps because of cultural traditions or a longer history with art vendors. The sale of A., standleyanum baskets is an important source of cash income, with both domestic and international sales. The destructive harvesting of palms threatens the industry and suggestions are made for sustainable harvest.     

Current utilization and further development of the palmyra palm (Borassus flabellifer L., Arecaceae) in Tamil Nadu state,India

The palmyra palm Borassus flabellifer,), a multipurpose tree of great utility, occurs extensively in Tamil Nadu state, India. Figuring in history, literature, and folklore of the state, it is exploited for food from the fruit and tuberous seedlings; beverage and sugar from the sap; fiber from the leaf and leafbase for brushes, cordage, weaving, and plaiting; trunk wood for construction and fuel; and numerous minor products. Increasing exploitation of the palmyra threatens the future supply of palm raw materials so important to rural populations. Integrated development of palmyra products for local and export markets, as well as management/conservation measures, are needed both to maximize the economic value of the products and to assure sustained yield from native stands.     

Miocene dispersal drives island radiations in the palm tribe Trachycarpeae (Arecaceae)

The study of three island groups of the palm tribe Trachycarpeae (Arecaceae/Palmae) permits both the analysis of each independent radiation and comparisons across the tribe to address general processes that drive island diversification. Phylogenetic relationships of Trachycarpeae were inferred from three plastid and three low-copy nuclear genes. The incongruent topological position of Brahea in CISP5 was hypothesized to be caused by a gene duplication event and was addressed using uninode coding. The resulting phylogenetic trees were well-resolved and the genera were all highly supported except for Johannesteijsmannia and Serenoa. Divergence time analysis estimated the stem of the tribe to be approximately 86 Ma and the crown to be 38 Ma, indicating that significant extinction may have occurred along this branch. Historical biogeographic analysis suggested that Trachycarpeae are of southern North American, Central American, or Caribbean origin and supports previous hypotheses of a Laurasian origin. The biogeography and disjunctions within the tribe were interpreted with respect to divergence times, the fossil record, and geological factors such as the formation of the Greater Antilles--Aves Ridge, the Bering and the North Atlantic land bridges, tectonic movement in Southeast Asia, climatic shifts between the Eocene and Pliocene, and volcanism in the Pacific basin. In considering the three major island radiations within Trachycarpeae, Miocene dispersal appears to have been the driving force in allopatric speciation and is highlighted here as an emerging pattern across the tree of life.     

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.     

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 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.     

Reproductive developmental complexity in the African oil palm (Elaeis guineensis, Arecaceae)

Species of the palm family (Arecaceae) are remarkably diverse in their inflorescence and floral morphologies, which make them a particularly interesting group for studies of reproductive development and its evolution. Using light and scanning electron microscopy, we describe inflorescence and flower development in the African oil palm Elaeis guineensis from the initiation of the inflorescence meristem to flower maturity. In mature palms, the inflorescence develops over 2-3 years and is characterized by individual stages within which differentiation may be either relatively slow, as in the case of early inflorescence meristem development, or rapid, as in the case of flower organogenesis. The female inflorescence bears floral triads composed of single pistillate flowers flanked by two abortive staminate flowers, whereas the male inflorescence contains single functional staminate flowers. This suggests a possible evolutionary movement from an ancestral hermaphrodite inflorescence form containing fully functional floral triads to the situation of temporal dioecy observed at present. Wild type flowers are compared to those bearing an epigenetic homeotic abnormality, known as mantled, involving an alteration of the identity of the organs in the fertile and sterile androecium.     

The cohune palm (Orbignya cohune,Arecaceae) in Belize: A survey of uses

Although common throughout Belize in lowland forests and agricultural systems, the cohune palm,Orbignya cohune is relatively little used compared with its historic importance, according to a 1992 survey discussed here. Among uses found to persist are leaves for thatch, fruits for oil, and palm heart as food. Exploitation required only simple technology and was most notable at socio-economic extremes. Areas from which cohunes were exploited rarely coincided with high-density stands of the species. Alternatives exist for all cohune products. It seems unlikely that this palm will be the basis of a large-scale industry in Belize, primarily because of the uncertainty of wild yields and the widespread availability of alternatives.     

Historical legacies in the geographical diversity patterns of New World palm (Arecaceae) subfamilies

The extent to which species richness patterns of the major palm subfamilies in the Americas are controlled by lineage history was studied. Based on the fossil record, we suggest that the subfamily Coryphoideae has followed a boreotropical dispersal route into Central and South America, whereas Calamoideae (tribe Lepidocaryeae), Ceroxyloideae and Arecoideae have Gondwana/South America-biased histories. However, Arecoideae has been present and diverse in both South and Central America at least since the early Tertiary. We used regression analyses to evaluate the relative importance of environmental factors and spatial variables (as substitutes for historical or other non-environmental factors) as determinants of geographical variation in species richness for each subfamily. Given the different lineage histories, we hypothesized that: (1) coryphoid richness should be least strongly controlled by the modern environment and exhibit a strong non-environmental bias towards Central and North America, reflecting its boreotropical invasion route, (2) calamoid species richness should exhibit a non-environmental bias towards South America, reflecting its long African–South American history, and (3) arecoid species richness should be most strongly environmentally determined, reflecting the long arecoid residency in both Central and South America. The regression analyses confirmed the hypothesized effects of lineage history on the geographical patterns in species richness. Hence, modern species richness patterns in the New World palm subfamilies strongly reflect their divergent biogeographical histories.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 113–125.     

Commonness of Amazonian palm (Arecaceae) species: Cross-scale links and potential determinants

The mechanisms that cause variation in commonness (abundances and range sizes) of species remain debated in ecology, and a repeatedly observed pattern is the positive relation between local abundances and larger scale range sizes. We used the Amazonian palm species (Arecaceae) to investigate the dependence between and potential determinants of commonness across three (local, landscape, continental) spatial scales. Commonness at the smaller scales (local abundance, landscape frequency) was estimated using data from 57 transects (5 × 500 m) in primary, non-inundated (terra firme) rainforest in a western Amazonian landscape, while commonness at the largest scale (continental range size) was estimated from digitized distribution maps. Landscape frequency was positively related to both local abundance and continental range size, which, however, were not related to each other. Landscape frequency was positively related to topographic niche breadth. Stem height correlated with continental range size and was the only species life-history trait related to any commonness measure. Distance from the study area to a species' range centre did not influence any of the commonness measures. The factors determining commonness in the Amazonian palm flora appear to be scale-dependent, with the unrelated local scale abundance and continental range size probably being controlled by different driving factors. Interestingly, commonness at the intermediate, landscape scale seems linked to both the smaller and the larger scale. Our results point towards topographic niche breadth at the smaller scales and stem height, possibly reflecting species' dispersal potential, at the continental scale as important determinants of commonness.     

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.