Reproductive patterns of selected understorey trees in the Malaysian rain forest: the apomictic species

HA, C. O., SANDS, V. E., SOEPADMO, E. & JONG, K., 1988. Reproductive patterns of selected understorey trees in the Malaysian raia forest: the apomictic species. Garcinia is the predominant representative of the Clusiaceae in the understorey of the lowland rain forest of Peninsular Malaysia. In Pasoh Forest Reserve the dioecious species G. paruifolia was investigated since no male trees, but only trees with structurally hermaphodite flowers which later set fruit, were found there. Pollination and embryological studies indicated a condition of non-pseudogamous agamospermy, with no viable pollen grains being formed in the staminodes, and the unfertilized egg cell giving rise to the embryo. Supportive studies were made of microspore development in G. forbesi , and of embryogenesis and seed development in G. malaccensis and G. scortechinii which also appear to be agamospermous. The role of either facultative or obligate apomixis in gene fixation in these dioecious species is discussed, and the possible adaptive significance of reproductive versatility among certain rain forest trees is examined.     

Studies in Garcinia, dioecious tropical forest trees: agamospermy

RICHARDS, A. J., 1990. Studies in Garcinia , dioecious tropical forest trees: agamospermy . In Garcinia , agamospermy is known or suspected for ten species. Most are facultative agamosperms, with males occurring, but males are probably lacking in G. mangostana . In G. mangostana and G. hombroniana , adventitious embryony occurs autonomously, and haploid parthenogenesis may also occur rarely. In G. parvifolia , it is reported previously that gametophytic agamospermy occurs, and adventitious embryos are suppressed. Autonomous endospermy is found in G. mangostana and G. parvifolia . In G. hombroniana , endosperm probably only develops after PEN fertilization. In G. hombroniana , some proembryos are formed precociously, and further proembryo formation is inhibited by sexual embryos. Asexual proembryos tend to occur in large ovules in small ovaries, and sexual embryos tend to occur in small ovules in large ovaries. It is considered that facultative agamospermy renders the genus Garcinia particularly suitable for the development of new types of fruit. Agamospermy in a dioecious genus, adventitious embryony in the absence of fertilization and/or pseudogamy, and the co-occurrence of gametophytic and sporophytic agamospermies in the same genus are unusual phenomena, which are discussed in the context of the lifestyle of a tropical forest tree. For G. hombroniana, n = 24, and for G. cowa, n = 26 are reported.     

Studies in Garcinia, dioecious tropical forest trees: the phenology, pollination biology and fertilization of G. hombroniana Pierre

RICHARDS, A. J., 1990. Studies in Garcinia , dioecious tropical forest trees: the phenology, pollination biology and fertilization of G. hombroniana Pierre . Garcinia hombroniana is a facultative agamosperm which is pollinated by Trigona bees. Nectar is restricted to the large discoid stigma (or pistillode in male flowers), which also captures and hydrates pollen. The 'wet' stigma and binucleate pollen suggest that Garcinia arose from hermaphrodite plants with a gametophytic self-incompatibility system.
On stigmas, nectar is secreted early on three or four successive days. On male pistillodes, nectar is secreted when anthers dehisce, on the second morning after anthesis. Pollen is most viable when freshly collected, but some viability remains four days after collection. Pollen germinates within 24 h of hydration. Similar results to pollinations are obtained by germinating pollen in 1 % sucrose.
Garcinia hombroniana flowers principally from January to June. Cultivated females are considered as 'big bang' strategists. Male flowers are considered as 'steady state' strategists.     

Spatial Scales of Pollen and Seed-Mediated Gene Flow in Tropical Rain Forest Trees

Gene flow via seed and pollen is a primary determinant of genetic and species diversity in plant communities at different spatial scales. This paper reviews studies of gene flow and population genetic structure in tropical rain forest trees and places them in ecological and biogeographic context. Although much pollination is among nearest neighbors, an increasing number of genetic studies report pollination ranging from 0.5–14 km for canopy tree species, resulting in extensive breeding areas in disturbed and undisturbed rain forest. Direct genetic measures of seed dispersal are still rare; however, studies of fine scale spatial genetic structure (SGS) indicate that the bulk of effective seed dispersal occurs at local scales, and we found no difference in SGS (Sp statistic) between temperate (N?=?24 species) and tropical forest trees (N?=?15). Our analysis did find significantly higher genetic differentiation in tropical trees (F _ST?=?0.177; N?=?42) than in temperate forest trees (F _ST?=?0.116; N?=?82). This may be due to the fact that tropical trees experience low but significant rates of self-fertilization and bi-parental inbreeding, whereas half of the temperate tree species in our survey are wind pollinated and are more strictly allogamous. Genetic drift may also be more pronounced in tropical trees due to the low population densities of most species.     

西双版纳热带森林树洞丰富度及其分配特点

树洞是森林生态系统的重要结构,对动物多样性的维持起着重要作用。为研究热带森林树洞的数量特征及其分配特点,选取西双版纳20hm2热带森林动态监测样地为研究对象,采用地面观测法调查了样地内所有胸径≥5cm活立木上的树洞。结果表明:(1)该样地森林的树洞密度为108.4个·hm-2,具有树洞的树1725株(占树木总数的6.22%),隶属于208个种;(2)树木个体的树洞数量随径级增加而增加,并在种间变化不均匀;(3)树木出现空心的概率随胸径级的增加而显著增加,并且在树种间存在显著差异;(4)与其他纬度地区森林的树洞密度比较,本研究结果支持了Boyle等(2008)提出的低纬度森林具有较高树洞密度的推测。     

MATING SYSTEM OF A TROPICAL RAIN FOREST TREE SPECIES

Canopy tree species are the dominant elements of the species-rich, fragile and endangered tropical rain forest ecosystems, yet little is known about the genetics of these species. We provide an estimate of the outcrossing rate in a population of Pithecellobium pedicellare, a large canopy tree in the tropical rain forests of Costa Rica. The outcrossing rate was high (t = 951 ± 0.021) and the pollen pool contributing to the progeny arrays used to measure outcrossing rate showed departure from homogeneity. The high outcrossing rate indicates that individuals scattered over a large area in this low density population could be bound with each other via outcrossing. However, despite a high outcrossing rate, the potential for inbreeding in this population is not negligible. We found a relatively large number of albino and chlorotic seedlings among the progeny arrays of several seed parents. Heterogeneity of pollen pool further indicates that the population, though widely outcrossed, may be structured. A more detailed study of mating systems in tropical rain forest trees may provide additional insights into the mating patterns of these trees. Such studies will be useful not only in understanding the dynamics of micro-evolutionary processes, but also in the conservation and management of tropical forest trees.     

Bird Assemblage and Visitation Pattern at Fruiting Elmerrillia tsiampaca (Magnoliaceae) Trees in Papua New Guinea

Most tropical trees produce fleshy fruits that attract frugivores that disperse their seeds. Early demography and distribution for these tree species depend on the effects of frugivores and their behavior. Anthropogenic changes that affect frugivore communities could ultimately result in changes in tree distribution and population demography. We studied the frugivore assemblage at 38 fruiting Elmerrillia tsiampaca, a rain forest canopy tree species in Papua New Guinea. Elmerrillia tsiampaca is an important resource for frugivorous birds at our study site because it produces abundant lipid-rich fruits at a time of low fruit availability. We classified avian frugivores into functional disperser groups and quantified visitation rates and behavior at trees during 56 canopy and 35 ground observation periods. We tested predictions derived from other studies of plant–frugivore interactions with this little-studied frugivore assemblage in an undisturbed rain forest. Elmerrillia tsiampaca fruits were consumed by 26 bird species, but most seeds were removed by eight species. The most important visitors (Columbidae, Paradisaeidae and Rhyticeros plicatus) were of a larger size than predicted based on diaspore size. Columbidae efficiently exploited the structurally protected fruit, which was inconsistent with other studies in New Guinea where structurally protected fruits were predominantly consumed by Paradisaeidae. Birds vulnerable to predation foraged for short time periods, consistent with the hypothesis that predator avoidance enhances seed dispersal. We identified seven functional disperser groups, indicating there is little redundancy in disperser groups among the regular and frequent visitors to this tropical rain forest tree species.     

Population structure, physiology and ecohydrological impacts of dioecious riparian tree species of western North America

The global water cycle is intimately linked to vegetation structure and function. Nowhere is this more apparent than in the arid west where riparian forests serve as ribbons of productivity in otherwise mostly unproductive landscapes. Dioecy is common among tree species that make up western North American riparian forests. There are intrinsic physiological differences between male and female dioecious riparian trees that may influence population structure (i.e., the ratio of male to female trees) and impact ecohydrology at large scales. In this paper, we review the current literature on sex ratio patterns and physiology of dioecious riparian tree species. Then develop a conceptual framework of the mechanisms that underlie population structure of dominant riparian tree species. Finally, we identify linkages between population structure and ecohydrological processes such as evapotranspiration and streamflow. A more thorough understanding of the mechanisms that underlie population structure of dominant riparian tree species will enable us to better predict global change impacts on vegetation and water cycling at multiple scales.     

Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.     

Fruit trees in a Malaysian rain forest

An inventory was made of 50 ha of primary lowland rain forest in Peninsular Malaysia, in which ca. 340,000 trees 1 cm dbh or larger were measured and identified to species. Out of a total plot tree flora of 820 species, 76 species are known to bear edible fruit. Especially diverse were the wild species of mango (Mangifera, Anacardiaceae, 12 spp.), mangosteen (Garcinia, Clusiaceae, 13 spp.), breadfruit (Artocarpus, Moraceae, 10 spp.) and rambutan (Nephelium, Sapindaceae, 5 spp.). Median population size for all species of fruit trees was 3.0 trees per ha and 0.2 adult trees per ha. Direct economic value of wild fruit trees was small; only one species has been very much collected and sold, Parkia speciosa (Fabaceae), amounting to less than US$20 per ha per year. The potential value of the species as genetic resources is very large: 24 species are cultivated, 38 edible species are congeneric with cultivated crops and at least 10 other species bear inedible fruit but are related to cultivated crops. We conclude that the Peninsular Malaysian rain forest is exceedingly rich in wild fruit trees, that these normally live at low densities, and that their principal economic value is as genetic resources.     

Long-distance pollen flow and tolerance to selfing in a neotropical tree species

Outcrossing rates, pollen dispersal and male mating success were assessed in Dicorynia guianensis Amshoff, a neotropical tree endemic to the Guiana shield. All adult trees within a continuous area of 40 ha (n = 157) were mapped, and were genotyped with six microsatellite loci. In addition, progenies were genotyped from 22 mature trees. At the population level, the species was mostly outcrossing (tm = 0.89) but there was marked variation among individuals. One tree exhibited mixed mating, confirming earlier results obtained with isozymes that D. guianensis can tolerate selfing. A Bayesian extension of the fractional paternity method was used for paternity analysis, and was compared with the neighbourhood method used widely for forest trees. Both methods indicated that pollen dispersal was only weakly related to distance between trees within the study area, and that the majority (62%) of pollen came from outside the study stand. Using maximum likelihood, male potential population size was estimated to be 1119, corresponding to a neighbourhood size of 560 hectares. Male mating success was, however, related to the diameter of the stem and to flowering intensity assessed visually. The mating behaviour of D. guianensis is a combination of long-distance pollen flow and occasional selfing. The species can still reproduce when it is extremely rare, either by selfing or by dispersing pollen at long distances. These results, together with the observation that male mating success was correlated with the size of the trees, could be implemented in management procedures aiming at regenerating the species.     

The behaviour of male orchid bees (Apidae,Hymenoptera, Insecta) and the question of leks

The male territories of two species of orchid bees, Eulaema meriana and Euglossa imperialis, are described. These territories consist of a perch, where the males display on the trunk of a tree, and a route flown from and back to the perch. Territories are located in treefalls or other large light gaps in tropical forest. The territories of these two species differ in the height of the perch and amount of light reaching it, size of the perch tree, and period of activity. A definition of lek is given. These male euglossines form facultative leks in large treefalls. The relationships between male territories, female foraging ranges, and collection of aromatic material by males are discussed.     

Intra‐specific relatedness,spatial clustering and reduced demographic performance in tropical rainforest trees

Intra‐specific negative density dependence promotes species coexistence by regulating population sizes. Patterns consistent with such density dependence are frequently reported in diverse tropical tree communities. Empirical evidence demonstrating whether intra‐specific variation is related to these patterns, however, is lacking. The present study addresses this important knowledge gap by genotyping all individuals of a tropical tree in a long‐term forest dynamics plot in tropical China. We show that related individuals are often spatially clustered, but having closely related neighbours reduces the growth performance of focal trees. We infer from the evidence that dispersal limitation and negative density dependence are operating simultaneously to impact the spatial distributions of genotypes in a natural population. Furthermore, dispersal limitation decreases local intra‐specific genetic diversity and increases negative density dependence thereby promoting niche differences and species coexistence as predicted by theory.     

Density‐dependence in common tree species in a tropical dry forest in Mudumalai,southern India

Abstract. Density‐dependence in tree population dynamics has seldom been examined in dry tropical forests. Using long‐term data from a large permanent plot, this study examined 16 common species in a dry tropical forest in southern India for density‐dependence. Employing quadrat‐based analyses, correlations of mortality, recruitment and population change with tree densities were examined. Mortality in 1–10 cm diameter trees was largely negatively correlated with conspecific density, whereas mortality in > 10 cm diameter trees was positively correlated. Mortality was, however, largely unaffected by the basal area and abundance of heterospecific trees. Recruitment was poor in most species, but in Lagerstroemia microcarpa (Lythraceae), Tectona grandis (Verbenaceae) and Cassia fistula (Fabaceae), species that recruited well, strong negative correlations of recruitment with conspecific basal area and abundance were found. In a few other species that could be tested, recruitment was again negatively correlated with conspecific density. In Lagerstroemia, recruitment was positively correlated with the basal area and abundance of heterospecific trees, but these correlations were non‐significant in other species. Similarly, although the rates of population change were negatively correlated with conspecific density they were positive when dry‐season ground fires occurred in the plot. Thus, the observed positive density‐dependence in large‐tree mortality and the negative density‐dependence in recruitment in many species were such that could potentially regulate tree populations. However, repeated fires influenced density‐dependence in the rates of population change in a way that could promote a few common species in the tree community.     

西双版纳青梅林的群落学研究

本文对分布在西双版纳勐腊县南部以龙脑香科植物版纳青梅为标志树种的热带森林作了群落学分析,认为它具有热带雨林的结构和基本特征,在性质上属于热带季节雨林。由于分布海拔偏高和生境特殊,它的上层乔木几乎常绿,在外貌上与望天树林和本地区典型的季节雨林有一定差异,在区系组成上向山地雨林过渡,它表现为一种季节雨林向山地雨林过渡的类型,同时也是一种热带北缘地区季节雨林的海拔极限类型。     

桂西南蚬木群落优势树种分布与环境因子的关系

蚬木(Excentrodendron hsienmu)是广泛分布于桂西南喀斯特山地的优良用材树种,了解影响蚬木群落物种分布的主要环境因子,对蚬木资源的有效保护具有重要意义。该研究基于桂西南蚬木群落的样地调查,测定了样地中乔木树种的重要值以及海拔、坡度、坡向、土壤养分等8个环境因子,采用Pearson相关分析研究了地形与土壤等环境因子间的相互关系,运用典范对应分析(CCA)方法对群落主要树种与环境因子间的关系进行了排序。结果表明:调查共记录到胸径≥1.0 cm、树高≥1.5 m的立木共176种,隶属于50科128属;群落乔木层以蚬木占绝对优势,主要伴生种有广西澄广花(Orophea anceps)、金丝李(Garcinia paucinervis)、割舌树(Walsura robusta)、苹婆(Sterculia nobilis)。相关分析显示海拔与土壤有机质、全氮间呈极显著正相关;除土壤pH之外,其余土壤肥力因子间均呈极显著正相关。CCA分析显示全钾、全磷对群落优势种的分布影响最为显著,坡向、坡度对优势种分布也具有重要作用。该研究结果揭示了影响蚬木群落物种分布的主要环境因子,为该区域的植被恢复措施提供了科学依据。     

Distinctive Phyllosphere Bacterial Communities in Tropical Trees

Recent work has suggested that in temperate and subtropical trees, leaf surface bacterial communities are distinctive to each individual tree species and dominated by Alpha- and Gammaproteobacteria. In order to understand how general this pattern is, we studied the phyllosphere bacterial community on leaves of six species of tropical trees at a rainforest arboretum in Malaysia. This represents the first detailed study of ‘true’ tropical lowland tree phyllosphere communities. Leaf surface DNA was extracted and pyrosequenced targeting the V1–V3 region of 16S rRNA gene. As was previously found in temperate and subtropical trees, each tree species had a distinctive bacterial community on its leaves, clustering separately from other tree species in an ordination analysis. Bacterial communities in the phyllosphere were unique to plant leaves in that very few operational taxonomic units (0.5%) co-occurred in the surrounding soil environment. A novel and distinctive aspect of tropical phyllosphere communities is that Acidobacteria were one of the most abundant phyla across all samples (on average, 17%), a pattern not previously recognized. Sequences belonging to Acidobacteria were classified into subgroups 1–6 among known 24 subdivisions, and subgroup 1 (84%) was the most abundant group, followed by subgroup 3 (15%). The high abundance of Acidobacteria on leaves of tropical trees indicates that there is a strong relationship between host plants and Acidobacteria in tropical rain forest, which needs to be investigated further. The similarity of phyllosphere bacterial communities amongst the tree species sampled shows a significant tendency to follow host plant phylogeny, with more similar communities on more closely related hosts.     

Low Inbreeding and High Pollen Dispersal Distances in Populations of Two Amazonian Forest Tree Species

Recent studies suggest that tropical tree species exhibit low inbreeding and high gene dispersal levels despite the typically low density of conspecifics in tropical forests. To examine this, we undertook a study of pollen gene dispersal and mating system of two Amazonian tree species. We analyzed 341 seeds from 33 trees at four microsatellite loci in a Carapa guianensis population from Brazil, and 212 seeds from 22 trees at four microsatellite loci in a Sextonia rubra population from French Guiana. Differentiation of allele frequencies among the pollen pool of individual trees was Φ_FT= 0.053 (95% CI: 0.027–0.074) for C. guianensis and Φ_FT= 0.064 (95% CI: 0.017–0.088) for S. rubra. The mean pollen dispersal distances were estimated at 69–355 m for C. guianensis , and 86–303 m for S. rubra , depending on the pollen dispersal model and the estimate of reproductive tree density used. The multi-locus outcrossing rate was estimated at 0.918 and 0.945, and the correlation of paternity at 0.089 and 0.096, for C. guianensis and S. rubra , respectively, while no significant levels of biparental inbreeding were detected. Comparing trees with high and low local density of conspecifics, we found no evidence for differences in inbreeding levels. The results are discussed within the framework of the emerging picture of the reproductive biology of tropical forest trees.     

Dual mycorrhizal colonization of forest-dominating tropical trees and the mycorrhizal status of non-dominant tree and liana species

The contribution of mycorrhizal associations to maintaining tree diversity patterns in tropical rain forests is poorly known. Many tropical monodominant trees form ectomycorrhizal (EM) associations, and there is evidence that the EM mutualism contributes to the maintenance of monodominance. It is assumed that most other tropical tree species form arbuscular mycorrhizal (AM) associations, and while many mycorrhizal surveys have been done, the mycorrhizal status of numerous tropical tree taxa remains undocumented. In this study, we tested the assumption that most tropical trees form AM associations by sampling root vouchers from tree and liana species in monodominant Dicymbe corymbosa forest and an adjacent mixed rain forest in Guyana. Roots were assessed for the presence/absence of AM and EM structures. Of the 142 species of trees and lianas surveyed, three tree species (the monodominant D. corymbosa, the grove-forming D. altsonii, and the non-dominant Aldina insignis) were EM, 137 were exclusively AM, and two were non-mycorrhizal. Both EM and AM structures were observed in D. corymbosa and D. altsonii. These results provide empirical data supporting the assumption that most tropical trees form AM associations for this region in the Guiana Shield and provide the first report of dual EM/AM colonization in Dicymbe species. Dual colonization of the Dicymbe species should be further explored to determine if this ability contributes to the establishment and maintenance of site dominance. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.