Seed size variation in the palm Euterpe edulis and the effects of seed predators on germination and seedling survival

Intraspecific variation in seed size is common in wild plant populations and has important consequences for the reproductive success of individual plants. Multiple, often conflicting evolutionary forces mediated by biotic as well as abiotic agents may maintain such a variation. In this paper we assessed seed size variation in a population of the threatened, commercially important palm Euterpe edulis in southeast Brazil. We investigated (i) how this variation affects the probability of attack by vertebrate and invertebrate post-dispersal seed predators, and (ii) if seed size influences the outcome of seeds damaged by beetles in terms of seed germination and early survival of seedlings. Euterpe edulis seeds varied in diameter from 8.3 to 14.1 mm. Neither insects nor rodents selected the seeds they preyed upon based on seed size. Seed germination and total, shoot and root biomasses of one-year seedlings were significantly and positively affected by seed size. Root biomass and seedling survival were negatively affected by seed damage caused by a scolytid beetle (Coccotrypes palmarum) whose adults bore into seeds to consume part of the endosperm, but do not oviposit on them. Seed size had a marginally significant effect on seedling survival. Therefore, if any advantage is accrued by E. edulis individuals producing large seeds, this is because of greater seed germination success and seedling vigor. If this is so, even a relatively narrow range of variation in seed size as observed in the E. edulis population studied may translate into differential success of individual plants.     

The history and mythology of Endymion nonscriptus

Background: Biological invasions are one of the major causes of biodiversity loss, yet remain rather understudied in tropical environments. The Australian palm tree Archontophoenix cunninghamiana was introduced into Brazil for ornamental purposes, but has become an invasive species in urban and suburban forest patches. The substitution of A. cunninghamiana by the native palm Euterpe edulis has been proposed as a management action.

Aims: We aimed to evaluate the regeneration potential of these two palm species in an Atlantic forest remnant in south-eastern Brazil where both species occur.

Methods: We compared seedling establishment and seed longevity of both species through seed sowing, and also measured the contribution of A. cunninghamiana to the local seed rain and seed bank.

Results: Nearly half of the non-anemochoric diaspores collected from the seed rain belonged to A. cunninghamiana, which represented a high propagule pressure in the community. The distribution of the alien palm seeds in the seed rain correlated with the distribution of nearby young and adult individuals inside the forest. Neither A. cunninghamiana nor E. edulis appeared to have a persistent seed bank in a burial experiment; seedling survival experiments suggested a much better performance for A. cunninghamiana, which had a survival rate of ca. 30% compared with a rate of only 3.5% for E. edulis.

Conclusions: The results suggest a higher regeneration capacity for the alien palm over the native species when co-occurring in a forest fragment. Management actions are thus proposed to reduce a potential biological invasion process.     

Effects of Local Plant Density and Incomplete Dichogamy on the Reproductive Success of the Rare Neotropical Palm Geonoma epetiolata

Density dependent processes are known to influence reproduction and establishment of plant populations. In this study, we evaluated the effects of local density and sexual expression on the reproductive success of the rare palm species Geonoma epetiolata in Costa Rica. We classified individuals in two density categories based on distances to the two nearest neighbors and recorded the occurrence of sexual overlap on each individual. Overlap between pistillate and staminate flowers in monoecious plants allows geitonogamous pollination, potentially reducing dependence on mates for reproduction. We measured plant size and light availability, and evaluated the influence of these variables on pollination success, fruit production, fruit abortion and seed mass. Pollination success significantly increased with density but there was no effect of sexual overlap. In contrast, there was no effect of density on the probability of initiated fruit, and a negative effect on fruit set. Fruit abortion was lowest in isolated plants with no sexual overlap. Plant size and light did not contribute to variation in reproductive success. In conclusion, the pollination advantage of plants in dense neighborhoods does not compensate reproductive losses incurred by fruit abortion, possibly due to mating among genetically related individuals. Geonoma epetiolata is threatened by habitat loss and poaching of seeds for the horticultural market. High fruit abortion rates associated with density in G. epetiolata suggest that seed collections from the remnant wild populations of this palm species may cause bottlenecks that further threaten population viability.     

Traditional Knowledge,Use, and Management of <Emphasis Type="Italic">Aphandra natalia</Emphasis> (Arecaceae) in Amazonian Peru

Traditional Knowledge, Use, and Management of Aphandra natalia (Arecaceae) in Amazonian Peru. Along the Pastaza and Urituyacu rivers in Peru, 100 informants in four villages knew 17 different uses of the piassaba fiber palm, Aphandra natalia (Balslev and Henderson) Barfod. Human consumption of fruits, extraction of fibers for trade, and harvesting leaves for thatch were the most frequently reported uses. Twenty uses were documented previously in the literature, and eight are reported here for the first time. Knowledge concerning the use of A. natalia among villagers varied significantly with ethnicity, village, and gender, but not with education or age; Amerindian informants knew more than non-Amerindian informants, and men knew more than women. Old and young informants knew about the same numbers of uses, and those who had little formal school training knew as many as those who had attended school for a longer time. Although 44% of our informants knew about fiber extraction, trade involving Aphandra fibers was not of great economic importance in our study area. Because fiber extraction has been documented as the most important use in all previous studies of A. natalia, it was surprising to find that people in one of the studied villages did not use A. natalia for that purpose at all. In contrast to previous observations, the extraction of fibers involved the destructive felling of the palms. We recommend implementing non-destructive extraction techniques so that local communities could exploit the fiber resource in a sustainable way. Non-destructive extraction methods involve cutting leaves from standing palms instead of felling the entire tree and leaving a few leaves on each extracted palm. However, given the limited economic incentive to the extractor, market mechanisms may be the greatest barrier to implementing sustainable extraction techniques.     

Demography of palm species in Brazil’s Atlantic forest: a comparison of harvested and unharvested species using matrix models

Surprisingly little is known about the effect of alternative harvesting methods and different means of reproduction on the population dynamics of plant species. Here, we test the hypothesis that habitat fragmentation leads to negative population growth rates for three palm species in Brazil’s biodiversity-rich and highly fragmented Atlantic forest. We compared the demography (λ and elasticities) of Astrocaryum aculeatissimum, Euterpe edulis and Geonoma schottiana in five fragments ranging in size from 19 to 3500 ha. A total of 4.05 ha were censused throughout the landscape studied. All individuals of the three palms were tagged in 2005 and their survival was monitored in 2006 and 2007. All new plants were also recorded. Summary matrices were constructed for each transition years by pooling data from all plots of all fragments together for each species. Based on summary matrices, A. aculeatissimum and G. schottiana asymptotic population growth rates were not significantly different from 1, suggesting that populations of these palms were stable. The projection of E. edulis populations was to decrease in size, with 95% confidence intervals of λ for these estimates failing to exceed one in the second transition year. A. aculeatissimum, E. edulis and G. schottiana survivorship was high for all stage classes and exceeded 85% for postseedling stages. We found that the more important vital rates were the survival rates in larger stages for all three species. Our results show that the maintenance of high levels of reproductive survivorship should be a focus of conservation strategies for these species, especially in the case of large E. edulis individuals.     

Palm wine harvesting by the Bassari threatens Borassus aethiopum populations in north-western Guinea

The use of the palm tree Borassus aethiopum Mart(Coryphoideae) by the Bassari ethnic group in the Republic of Guinea wasstudied. Ethnic groups have multiple uses for B. aethiopumthroughout its distribution in tropical Africa. The Bassari use this palm mostlyas a palm wine resource. As their harvesting technique leads to the death of thepalm, the populations of this species are highly endangered in the areapopulated by the Bassari. The fast decline of an important resource has recentlygained attention at governmental level. In order to conserve the populations ofthis palm, it is essential to understand whether the Bassari use a lethaltechnique because of socio-economic reasons or due to a lack of knowledge. Thepresent study aims to answer this question. We collected the data in thenorth-western part of the Republic of Guinea in May 1996 by field observationsand by using 'Rapid Rural Appraisal' tools consisting of interactiveinterviews with palm wine harvesters, collectors, carriers and sellers. Afterconstructing a woody ladder fixed to the stem the Bassari extract the sap bycutting a hole into the palm heart and by removing a small part of the palmheart two times per day. Individual trees produce on average 10 lper day. After 35–45 days the sap production ceases and thepalm dies. A harvester kills on average 56 individuals of B.aethiopum per year. As a consequence of this lethal techniqueexploiters have to constantly move through and between palm populations. Thereis no reforestation of the degraded areas. The palm wine extraction ofB. aethiopum is the main activity of the Bassari for 7months of the year. For harvesters, collectors, carriers andsellers, palm wine extraction provides one of the most important annual incomes.The harvester pays a monthly tax of USD 5 to the forestry administration.Despite its economic importance, the Bassari palm wine harvesters rangeB. aethiopum as the sixth most important plant species in the area. The paradoxical situation of acommunity that threatens a resource that it is highly dependent on may beexplained by the following circumstances. Firstly, the exploiters seek a maximumprofit on a short term basis. Secondly, the exploiters apparently do not takeinto consideration the total value of other potential products from theBorassus. Thirdly, the Bassari do not have an importanttradition for exploiting B. aethiopum so that there is noapparent cultural alliance between the Bassari and the palm. Finally, the rulesand regulations for harvesting Borassus products favourquick profit. The extinction of the palm population studied can be expectedwithin about 25 years if no protection measure is taken.     

Quantification and characterisation of <Emphasis Type="Italic">Trichoderma</Emphasis> spp. from different ecosystems

Basal stem rot of oil palm caused by Ganoderma boninense is of major economic importance. Observations of the low incidence of disease due to Ganoderma species in natural stands, suggest that the disease is kept under control by some biological means. Trichoderma spp. are saprophytic fungi with high antagonistic activities against soil-borne pathogens. However, their abundance and distribution are soil and crop specific. Trichoderma species have been found to be concentrated in the A1 (0–30 cm) and Be soil horizons (30–60 cm), although the abundance of Trichoderma was not significantly different between the oil palm and non-oil palm ecosystems. Characterisation of Trichoderma isolates based on cultural, morphological and DNA polymorphism showed that T. harzianum, T. virens, T. koningii and T. longibrachiatum made up 72, 14, 10 and 4% of the total Trichoderma isolates isolated. As Trichoderma species are present in the oil palm ecosystem, but at lower numbers and in locations different from those desired, soil augmentation with antagonistic Trichoderma spp. can be developed as a strategy towards integrated management of basal stem rot of oil palm.     

Superior quality agar from <Emphasis Type="Italic">Gracilaria</Emphasis> species (Gracilariales,Rhodophyta) collected from the Gulf of Mannar,India

Gracilaria edulis, G. crassa, G. foliifera, and G. corticata are naturally occurring agarophytes of Indian waters. These agarophytes were evaluated for their agar contents using an improved process recently reported by us (US Patent 2005/0267296A1). The effect of different concentrations of NaOH in the alkali treatment was studied for optimizing the extraction conditions. These Gracilaria species of Indian waters produced agars, both native and alkali treated, with different properties confirming the heterogeneity of the agar polymers in this genera, as one would expect. Among these, G. edulis and G. crassa produced agar polymers having high gel strengths of 490 ± 8.16 and 800 ± 15.4 g cm⁻², respectively, with 8% NaOH treatment as opposed the low gel strength agars that have been reported in the literature to date.     

Enhancing biological control of basal stem rot disease (<Emphasis Type="Italic">Ganoderma boninense</Emphasis>) in oil palm plantations

Basal Stem Rot (BSR) disease caused by Ganoderma boninense is the most destructive disease in oil palm, especially in Indonesia and Malaysia. The available control measures for BSR disease such as cultural practices and mechanical and chemical treatment have not proved satisfactory due to the fact that Ganoderma has various resting stages such as melanised mycelium, basidiospores and pseudosclerotia. Alternative control measures to overcome the Ganoderma problem are focused on the use of biological control agents and planting resistant material. Present studies conducted at Indonesian Oil Palm Research Institute (IOPRI) are focused on enhancing the use of biological control agents for Ganoderma. These activities include screening biological agents from the oil palm rhizosphere in order to evaluate their effectiveness as biological agents in glasshouse and field trials, testing their antagonistic activities in large scale experiments and eradicating potential disease inoculum with biological agents. Several promising biological agents have been isolated, mainly Trichoderma harzianum, T. viride, Gliocladium viride, Pseudomonas fluorescens, and Bacillus sp. A glasshouse and field trial for Ganoderma control indicated that treatment with T. harzianum and G. viride was superior to Bacillus sp. A large scale trial showed that the disease incidence was lower in a field treated with biological agents than in untreated fields. In a short term programme, research activities at IOPRI are currently focusing on selecting fungi that can completely degrade plant material in order to eradicate inoculum. Digging holes around the palm bole and adding empty fruit bunches have been investigated as ways to stimulate biological agents.     

Staggered Flowering in Four Sympatric Varieties of Geonoma cuneata (Palmae)1

Phenology, flowering biology, and pollination were studied for one year in four sympatric varieties of the polymorphic palm Geonoma cuneata in Ecuador. Flowering seasons of the three most common varieties were significantly staggered resulting in minimal temporal overlap. No marked differences were found with respect to flowering biology or pollination. The implication of these findings for explaining the complicated morphological variation pattern found in G. cuneata and similar species complexes is discussed.     

Topographic and spatial controls of palm species distributions in a montane rain forest, southern Ecuador

The northern Andes harbour a flora that is as species-rich or even richer than the 18-times larger lowland Amazon basin. Gaining an understanding of how the high species richness of the Andean region is generated and maintained is therefore of particular interest. Environmental sorting due to elevational gradients in climate has been emphasized as a driver of vegetation distribution and plant community assembly in tropical mountain areas such as the Andes for two centuries, while alternative mechanisms have been little studied. Here, we investigated the importance of topography and spatial location as factors controlling species distributions in a palm community in a montane rain forest landscape in the Andes of southern Ecuador (1900–2150 m above sea level). Eleven species were present: Aiphanes verrucosa, Ceroxylon parvifrons, Chamaedorea pinnatifrons, Dictyocaryum lamarckianum, Euterpe precatoria, Geonoma densa, Geonoma orbignyana, Geonoma paradoxa, Prestoea acuminata and Wettinia aequatorialis. To study their spatial distribution, forty 250 m² (5 × 50 m²) plots were laid out perpendicular to four paths that were categorized into three areas and two topographic units (ridges and gullies). Mantel tests and indicator species analysis showed that both topography and spatial location imposed strong controls on palm species distributions at the study site. Our results suggest that species distributions in the studied montane forest landscape were partly determined by the species’ habitat requirements, but also by unknown spatial effects. Although a number of possible explanations exist for the latter, such as unmeasured environmental variables and historical disturbance events, we believe dispersal limitation is likely to be involved. Furthermore, although the gully- or ridge-association of some species corresponded to their general elevational ranges in southern Ecuador, this was not the case for other species. Based on such considerations, we conclude that elevational climatic gradients are likely to only form part of the explanation for the topographic effects on palm species distributions at the study site. Other factors must also be involved, notably wind-exposure and hydrology, as discussed for lowland palm communities. Our results show that to understand plant community assembly in the tropical montane forests of the Andes it is too simple to focus just on environmental sorting by elevational climatic gradients.     

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.     

Two new species and two new combinations in Mesoamerican Ericaceae

Satyria ventricosa andVaccinium monteverdense from Panama and Costa Rica, respectively, are described and illustrated.Gonocalyx megabracteolatus andVaccinium talamancense are new combinations. Relationships of the new species are discussed. Keys are provided for the entire genusGonocalyx, and forSatyria in Mesoamerica.

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Resumen  Se describenSatyria ventricosa yVaccinium monteverdense de Panamá y Costa Rica, respectivamente, con sus ilustraciones. Se hace una nueva combinación deGonocalyx megabracteolatus yVaccinium talamancense. También se discuten las relaciones entre las nuevas especies. Se incluyen claves para el género completoGonocalyx y para el géneroSatyria en Mesoamérica.

     

A new species of Columnea and range extension in the Gesneriaceae from Costa Rica

Columnea lariensis is described and illustrated as a new species from the Talamanca Mountains, Costa Rica. It is similar to C. canarina from Panama. Alloplectus weirii is here reported for the first time from Mesoamerica and an illustration is provided.

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Resumen   Columnea lariensis de la Cordillera de Talamanca, Costa Rica, es descrita e ilustrada. Es similar a C. canarina de Panamá. Alloplectus weirii se reporta para Mesoamérica por primera vez y se incluye una ilustración de la misma.

     

Dinitrogen-fixation by three neotropical agroforestry tree species under semi-controlled field conditions

Cultivating dinitrogen-fixing legume trees with crops in agroforestry is a relatively common N management practice in the Neotropics. The objective of this study was to assess the N₂ fixation potential of three important Neotropical agroforestry tree species, Erythrina poeppigiana, Erythrina fusca, and Inga edulis, under semi-controlled field conditions. The study was conducted in the humid tropical climate of the Caribbean coastal plain of Costa Rica. In 2002, seedlings of I. edulis and Vochysia guatemalensis were planted in one-meter-deep open-ended plastic cylinders buried in soil within hedgerows of the same species. Overall tree spacing was 1 × 4 m to simulate a typical alley-cropping design. The ¹⁵N was applied as (NH₄)₂SO₄ at 10% ¹⁵N atom excess 15 days after planting at the rate of 20 kg [N] ha⁻¹. In 2003, seedlings of E. poeppigiana, E. fusca, and V. guatemalensis were planted in the same field using the existing cylinders. The ¹⁵N application was repeated at the rate of 20 kg [N] ha⁻¹ 15 days after planting and 10 kg [N] ha⁻¹ was added three months after planting. Trees were harvested 9 months after planting in both years. The ¹⁵N content of leaves, branches, stems, and roots was determined by mass spectrometry. The percentage of atmospheric N fixed out of total N (%N_f) was calculated based on ¹⁵N atom excess in leaves or total biomass. The difference between the two calculation methods was insignificant for all species. Sixty percent of I. edulis trees fixed N₂; %N_f was 57% for the N₂-fixing trees. Biomass production and N yield were similar in N₂-fixing and non-N₂-fixing I. edulis. No obvious cause was found for why not all I. edulis trees fixed N₂. All E. poeppigiana and E. fusca trees fixed N₂; %N_f was ca. 59% and 64%, respectively. These data were extrapolated to typical agroforestry systems using published data on N recycling by the studied species. Inga edulis may recycle ca. 100 kg ha⁻¹ a⁻¹ of N fixed from atmosphere to soil if only 60% of trees fix N₂, E. poeppigiana 60–160 kg ha⁻¹ a⁻¹, and E. fusca ca. 80 kg ha⁻¹ a⁻¹.     

Flowering biology ofGeonoma irena andG. cuneata var.sodiroi (Arecaceae)

The phenology and flowering of two sympatric understory palms from western Ecuador, until recently regarded as conspecific, is described.Geonoma irena Borchs. flowers throughout the year. Anthesis lasts 11–14 weeks per inflorescence, with overlapping male and female phases. Flowers open in the midmorning, and are visited mainly by meliponid and halictid bees.Geonoma cuneata var.sodiroi (Burret)Skov flowers from December to April, with a distinct peak in Feburary. Anthesis lasts 6–8 days per inflorescence, with non-overlapping male and female phases. Flowers open at dawn, and are visited mainly by drosophilid and sphaerocerid flies. The flowering pattern of the two species match different specific behavioural features of their insect visitors. The study provides an example of how differences in reproductive biology may act as an important barrier to gene flow between related, co-occurring taxa.     

A profile of palm heart extraction in the Amazon estuary

Many of the processed palm hearts consumed throughout the world are derived from the açaípalm (Euterpe oleraceaMart.)), which grows abundantly in floodplain forests of the Amazon estuary. Palm heart extraction began in the estuary in the 1970s and there are now hundreds of canning factories and some 50 distribution firms in this region. Annual profits of the canning factories range from $30,000 to $50,000 while profits for distribution firms frequently exceed $500,000/year. But there are several indications that this economic boon will be short-lived: factory closings are frequent, palm hearts are much smaller now than in the past, and mortality of palm trees is high in stands subjected to frequent palm heart harvest. However, the açaípalm is well suited for management because of its abundance, rapid growth, and multistemmed life form. Under management, palm hearts can be harvested from the same clump over many years through controlled thinning. The management of açaístands could result in significant long-term savings for palm heart factories. Indeed, açaímanagement may offer one of the best opportunities to date for sustained use of some Amazonian forests.     

Three new species in the neotropical genus <Emphasis Type="Italic">Clidemia</Emphasis> (Melastomataceae: Miconieae)

We here propose and describe three new species of Clidemia: C. aguilarii from Costa Rica and Panama; C. aurantiaca from Costa Rica, Panama, and Ecuador; and C. subpeltata, a Costa Rican endemic. Phenological notes, distribution maps, line drawings, color images of the live plants in the wild, and discussions comparing the new species to their presumed closest relatives are provided.     

Two new species of Blechnum (Blechnaceae) from the neotropics

Blechnum moranianum is a new species similar toB. loxense, andB. nigrum is segregated from theB. fragile complex. Both are described and illustrated as result of the author's taxonomic fern work in Costa Rica. Blechnum moranianum es una especie nueva similar aB. loxense, yB. nigrum es segregada del complejoB. fragile. Ambas son aquí descritas e ilustradas como resultado de los trabajos taxonómicos del autor en los helechos de Costa Rica.     

Identification of <Emphasis Type="Italic">Ganoderma</Emphasis>, the causal agent of basal stem rot disease in oil palm using a molecular method

From comparison of the alignments of the internally transcribed spacers (ITS) of ribosomal DNA from Ganoderma associated with oil palm basal stem rot (BSR) and other Ganoderma species, two specific primer pairs were selected to provide a specific DNA amplification of pathogenic Ganoderma in oil palm. Each primer pair produced a single PCR product of about 450 bp (for primer pair IT1–IT2) and 334 bp (for primer pair IT1–IT3) when oil palm Ganoderma DNA was used. No PCR amplification product was observed when other Ganoderma species DNA was used in PCR amplification with these primer pairs. Three specific restriction enzyme sites were identified in the ITS and intergenic spacer (IGS1) regions. The restriction enzymes MluI, SacI and HinfI were used to digest the ITS-PCR product and restriction enzymes TfiI, ScaI and HincII were used to digest the IGS1-PCR product. Of the three restriction enzymes used in each rDNA region, MluI specifically digested the ITS regions, and TfiI specifically digested the IGS1 region of oil palm Ganoderma. Analysis of the published ITS nucleotide sequences of 31 Ganoderma species showed that the MluI restriction site was not present in other Ganoderma species. The use of both specific primers and restriction enzyme analysis can be applied as a standard protocol to identify pathogenic Ganoderma in oil palm. In this study, the use of specific primers and PCR-RFLP analyses of the rDNA gave consistent results for the characterisation of pathogenic Ganoderma, and indicated that Ganoderma strains associated with BSR disease in oil palms belong to a single species.