Influence of Forest Type and Tree Species on Canopy‐Dwelling Beetles in Budongo Forest,Uganda1

Beetles were collected on 64 trees of four species (Cynometra alexandri C. H. Wright, Rinorea beniemis (Welwitsch ex Olivier) Kuntze, Teclea nobilis Delile, and Trichilia rubescens Olivier) in Budongo Forest, Uganda, using an insecticidal fogging technique. Selected tree species were abundant, taxonomically not closely related, and different in the shape of leaves, growth form, and size, with heights between 7 and 35 m. Trees were fogged in an old primary forest stand, in an area of secondary forest where selective logging was performed, and in a swamp forest. Eight conspecific trees per forest type were fogged. A total of 29,736 beetles were collected from all trees that could be assigned to 1433 (morpho)‐species; 41.6 percent were singletons and 89.6 percent of species were found with less than ten individuals. Abundant beetle taxa included Latridiidae (N= 4093), Chrysomelidae (3952), Staphylinidae (2931), Apioninae (2621), and Curculionidae (2457). Most species‐rich groups were Staphylinidae (N= 196 spp.), Curculionidae (189), and Chrysomelidae (148). Abundance increased in the order: primary < secondary < swamp forest. Due to the relatively high dominance of some species in the secondary forest, species richness increased in the order: secondary < primary < swamp forest. Beta diversity measures and factor analysis showed distinct differences among forest types but higher similarity of beetle communities on different tree species within one forest type. The taxonomic distribution of beetles in the secondary forest was more heterogeneous than in the primary forest. Analyses of the data revealed low host specificity even for phytophagous beetles, underlining the importance of habitat structure and chance effects on the spatial distribution of beetles in the canopy of Budongo Forest.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

Arthropod Abundance,Canopy Structure,and Microclimate in a Bornean Lowland Tropical Rain Forest1

This study applies a novel, vertically stratified fogging protocol to document arthropod abundance, density, and biomass across a vertical gradient in a primary, lowland dipterocarp forest canopy in Borneo. We fogged arthropods at 5 m vertical intervals and 20 m horizontal intervals along six full‐canopy transects and measured leaf surface areas along the same transects. The results show that arthropod biomass in the aboveground regions was 23.6 kg/ha, the abundance was 23.9 million individuals/ha, and the density on leaf surfaces was 280 individuals/m² leaf area. All three numbers are five to ten times higher than estimated by previous surveys of tropical lowland rain forest canopies using mass‐collection techniques. Arthropod abundance and biomass were analyzed in relation to canopy structure, composition, vapor pressure deficit (VPD), photosynthetic photon flux density (PPFD), and height. Using stepwise regression we found that 13 of 14 arthropod groups had significant positive relationships with one‐sided leaf area, 11 had significant negative relationships with VPD, 3 had significant relationships with height, and none showed positive relationships with light. Classifying the 14 taxa based on their responses to leaf area and VPD created three groups that corresponded roughly to the biology of these taxa. This study suggests that the biomass and abundance, and perhaps therefore—by extrapolation—the biodiversity, of tropical canopy arthropods may be very much higher than previously estimated.     

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.     

Variation in Seedling Density and Seed Predation Indicators for the Emergent Tree Dipteryx panamensis in Continuous and Fragmented Rain Forest1

Seedling density and the condition of stony endocarps of the tree Dipteryx panamensis were assessed in protected continuous forest and two forest fragments exposed to hunting and selective logging. Seedling density was higher in forest fragments than in continuous forest, while more whole endocarps and fewer chewed and half endocarps were found in fragments, indicating lower seed predation at fragment sites. These findings appear to contradict two earlier D. panamensis studies and we discuss methodological differences that could account for our disparate results. Hunting and fragmentation effects on mammal populations are suggested as a cause for the altered recruitment pattern in fragments.     

Home Range Size of Sympatric Squirrel Species Inhabiting a Lowland Dipterocarp Forest in Malaysia1

Home range sizes and spatial overlap of four sympatric squirrel species were investigated in a lowland dipterocarp forest in Malaysia using a radio‐tracking method. The population density of Callosciurus caniceps was highest and C. notatus was next highest, while C. nigrovittatus and Lariscus insignis were scarce. C. caniceps was larger than C. nigrovittatus and C. notatus while L. insignis was extremely small. For females, home range size was smaller in L. insignis than Callosciurus spp., which may support the body weight hypothesis: larger species have larger home ranges. Among the three Callosciurus species, female C. caniceps had the smallest home range. These differences were accounted for by habitat characteristics rather than by density or body weight; C. caniceps was dominant in bushy areas and used crowded small trees while C. notatus and C. nigrovittatus used large trees in the forest. In this study, home range size did not change seasonally; this differs from studies in temperate regions, possibly because food availability is much less variable among seasons in tropical rain forest. Home range overlap among heterospecific individuals was common but different species seemed to partition space by using different vertical levels of the forest. Consequently, the home range size and spatial overlap of sympatric squirrel species may be affected by habitat diversity in tropical rain forest.     

Fine‐scale Spatial Genetic Structure of Ten Dipterocarp Tree Species in a Bornean Rain Forest

Fine‐scale spatial genetic structure is increasingly recognized as an important factor in the studies of tropical forest trees as it influences genetic diversity of local populations. The biologic mechanisms that generate fine‐scale spatial genetic structure are not fully understood. We studied fine‐scale spatial genetic structure in ten coexisting dipterocarp tree species in a Bornean rain forest using microsatellite markers. Six of the ten species showed statistically significant fine‐scale spatial genetic structure. Fine‐scale spatial genetic structure was stronger at smaller spatial scales (≤ 100 m) than at larger spatial scales (> 100 m) for each species. Multiple regression analysis suggested that seed dispersal distance was important at the smaller spatial scale. At the larger scale (> 100 m) and over the entire sample range (0–1000 m), pollinators and spatial distribution of adult trees were more important determinants of fine‐scale spatial genetic structure. Fine‐scale spatial genetic structure was stronger in species pollinated by less mobile small beetles than in species pollinated by the more mobile giant honeybee (Apis dorsata). It was also stronger in species where adult tree distributions were more clumped. The hypothesized mechanisms underlying the negative correlation between clump size and fine‐scale spatial genetic structure were a large overlap among seed shadows and genetic drift within clumped species.     

The Potential of Tree Rings for the Study of Forest Succession in Southern Mexico

Studies of tropical secondary forest succession face strong limitations due to the slow pace of succession and the time-consuming task of monitoring processes. The occurrence of tree rings in secondary forest trees may help expand our knowledge on succession in these systems and may be useful for fallow dating in chronosequence studies. We examine here the potential of tree rings to study forest succession by sampling 70 species along chronosequences of dry and wet forests in southern Mexico. Based on wood anatomical features, we estimated that about 37 percent of the species presented distinct growth rings useful for ring studies. Overall, maximum number of rings matched well the interview-based fallow ages but, at some sites, trees had consistently higher numbers of rings, probably due to errors in fallow ages derived from interviews. Best fallow age estimations were obtained by examining rings in both pioneer and nonpioneer species. Reconstruction of species' establishment dates revealed that pioneer and nonpioneer species establish early during succession, and that species of both groups continue to recruit after many years. Our study clearly shows that tree ring analysis is a promising tool for studies on secondary forest succession in the tropics.     

Rapid Recovery of Biomass, Species Richness, and Species Composition in a Forest Chronosequence in Northeastern Costa Rica

Secondary forests are a vital part of the tropical landscape, and their worldwide extent and importance continues to increase. Here, we present the largest chronosequence data set on forest succession in the wet tropics that includes both secondary and old-growth sites. We performed 0.1 ha vegetation inventories in 30 sites in northeastern Costa Rica, including seven old-growth forests and 23 secondary forests on former pastures, ranging from 10 to 42 yr. The secondary forest sites were formerly pasture for intervals of <1–25 yr. Aboveground biomass in secondary forests recovered rapidly, with sites already exhibiting values comparable to old growth after 21–30 yr, and biomass accumulation was not impacted by the length of time that a site was in pasture. Species richness reached old-growth levels in as little as 30 yr, although sites that were in pasture for > 10 yr had significantly lower species richness. Forest cover near the sites at the time of forest establishment did not significantly impact biomass or species richness, and the species composition of older secondary forest sites (>30 yr) converged with that of old growth. These results emphasize the resilience of tropical ecosystems in this region and the high conservation value of secondary forests.
     

热带森林树种多度和丰富度空间分布特征研究

用巴拿马50 hm2森林动态监测样地内直径≥1 cm的树种资料,分析了该样地树种多度(个体数)和丰富度(物种数)及其方差和变异系数在6个取样尺度(5 m×5 m,10 m×10 m,20 m×20 m,25 m×25 m,50 m×50 m,100 m×100 m)的变化规律.结果显示:(1)由于多度的可加性,不同取样尺度在样地内树种多度的变化表现出一致性;随取样尺度的增加,多度方差呈线性增加,而变异系数呈线性减小.(2)丰富度随取样尺度的变化较为复杂,随取样尺度的增加,丰富度方差呈非线性变化,在取样尺度为25 m×25 m时方差最大;变异系数随取样尺度的增加而呈线性减小.研究表明,大尺度的多度值可以由小尺度的多度值通过外推法估计,而丰富度却不能,在生物多样性的保护和管理中不能简单地从一个取样尺度的生物丰富度推测另一个取样尺度丰富度.     

Fine partitioning of epiphyte habitat within Johansson zones in tropical Australian rain forest trees

For over three decades, the Johansson zones have been widely used in epiphyte studies as a way of stratifying the host tree into habitat zones. The usefulness of this system, however, has been questioned. We test the effectiveness of the Johansson zones by grouping epiphyte species by the substrate and microclimatic attributes of their individual occurrences and assessing the fidelity of these groups to the Johansson zones. Habitat characteristics were recorded for every individual epiphyte on 30 trees in the lower montane rain forests of northeastern Australia. Twenty‐four epiphyte species were agglomerated into four groups using Ward's method. Group 4 was highly distinct and included shade‐loving species and nomadic vines from the lower zones of the host trees. Group 3 contained species from the most exposed habitats. Group 1 had higher light levels and lower substrate thickness than Group 2, yet both groups had close to identical distributions over the Johansson zones. This suggests that groups of epiphyte species may utilize different micro‐sites within the same zone. While the Johansson zones are a useful tool in epiphyte studies, finer partitioning of habitat within the host tree may be missed.     

海南霸王岭热带山地雨林森林循环与树种多样性动态

通过对海南岛霸王岭热带山地雨林的调查 ,研究了热带山地雨林树种多样性特征随森林循环的动态变化规律。结果表明 :( 1 )热带山地雨林森林循环不同阶段斑块在森林景观中所占的面积比例分别是 :林隙阶段 ( G)占 38.5 0 % ,建立阶段 ( B)占 2 8.5 0 % ,成熟阶段 ( M)占 2 7.0 0 % ,衰退阶段 ( D)占 6 .0 0 %。 ( 2 )热带山地雨林中乔木树种的密度随森林循环的变化趋势是由 G→B→M呈现出逐渐增加的趋势 ,以成熟阶段达到最大 ,而到衰退阶段又趋于下降。灌木树种则表现出 G阶段斑块的密度最大 ,B阶段的最小 ,从 B到 M有所增加 ,到 D又稍有下降。 ( 3)热带山地雨林中不同高度级和不同径级的树木的密度在森林循环的不同阶段表现出不同的增减趋势 ,其随森林循环过程呈现出的动态变化可能与不同阶段斑块内的空间、环境及物种生物学特性有关。 ( 4 )热带山地雨林中树木的平均胸径、平均高、平均胸高断面积、平均单株材积随森林循环过程呈现出不断增加的趋势 ,其中平均胸径和平均高随森林循环的变化较为平缓 ,而平均胸高断面积和平均单株材积之变化较为陡急。 ( 5 )热带山地雨林森林循环不同阶段的物种多样性指数不同 ,其中 G和 B阶段的物种丰富度和多样性指数值较接近 ,M阶段的物种丰富度达到最大 ,D阶段则最小。     

Edaphic and Floristic Variation within a 1‐ha Plot of Lowland Amazonian Rain Forest1

Several studies in lowland tropical rain forests have documented effects of local‐scale topographic variation on plant species distribution and abundance patterns. Few studies have compared the distribution patterns of more than one plant group, however, and even fewer have related these to measured physical and chemical soil characteristics. Here, we document such soil characteristics within a square 1‐ha plot in Amazonian Ecuador, and compare them to the distribution patterns of terrestrial pteridophytes, angiosperm ground herbs, and palms. Substantial variation in soil properties was found within the plot. The three plant groups showed highly correlated floristic patterns within the 1‐ha plot even after the effect of geographical distances had been taken into account. Mantel tests yielded significant correlations between edaphic patterns, as measured by distances in various soil and topographic characteristics, and floristic patterns. For all three plant groups, differences in elevation within the plot were highly correlated with floristic distances, and for terrestrial pteridophytes and palms, distances in soil calcium content and sand content were also important. Our results resembled those obtained at wider spatial scales with the same plant groups, which indicates that soil factors may play an important role for distribution and beta diversity of plants, even at the local scale.     

夜间低温对2种热带雨林树种幼苗叶绿素荧光的影响

研究了夜间低温(5℃±1℃,12h,连续3个晚上)对生长于3种光照条件下(100%、25%和8%光照)的热带雨林冠层树种绒毛番龙眼(Pometiatomentosa)和中层树种滇南风吹楠(Horsfieldiatetratepala)幼苗叶绿素荧光的影响。结果表明:低温处理使生长于全光下的绒毛番龙眼幼苗的Fv/Fm(PSII最大光化学量子产量)急剧降低,中午发生了强烈的光抑制,但随着低温胁迫的解除Fv/Fm能很快恢复,表明没有发生不可逆的光氧化损伤。低温使生长于全光和25%光照条件下2个树种幼苗的NPQ(非光化学猝灭,热耗散)受到抑制,但没有引起2个树种幼苗Fo(初始荧光)的升高,不会导致幼苗PSII反应中心的失活。2种热带雨林植物对低温的抗性可能与这些植物的地理分布和历史渊源有关。     

Ants Induce Domatia in a Rain Forest Tree (Vochysia vismiaefolia)1

In Amazonian rain forest trees of Vochysia vismiaefolia (Vochysiaceae), ants were found to induce twig structures that resembled classical ant domatia. This phenomenon is novel for ant‐plants, which commonly develop domatia without the activity of ants. Eight species of ants were recorded inside the domatia of six individual trees, but domatia were most numerous and characteristic when induced and inhabited by an undescribed species of Pseudomyrmex on two trees. To investigate the mechanism of domatium growth, we drilled holes into young twigs: the expansion of the twig diameter surrounding the holes was significantly accelerated, comparable to domatia formation. Domatia induction is discussed as a putative step in the evolution of ant‐plants.     

Effects of Canopy Gaps,Topography, and Soils on the Distribution of Woody Species in a Central Brazilian Deciduous Dry Forest1

The interrelationships between the distribution of woody species and environmental variables were investigated in an area of deciduous dry forest in Santa Vitöria, central Brazil. This is the first study of a vanishing type of dry forest which grows on base-rich soils originating from the basalt bedrocks of southern Goiás and western Minas Gerais. A survey of topography, soil properties, canopy gaps and woody plants (≥5 cm diameter at the base of the stem) was conducted in 50–15 × 15 m quadrats. The soils were classified into the following soil series: Hapludolls → Haplustolls → Haplustolls → Ustropepts → Rhodustalfs. This series corresponded to a gradient of increasing elevation and effective soil depth and decreasing slope gradient, soil organic matter and total exchangeable bases. A canonical correspondence analysis and a detrended correspondence analysis indicated that plant species’abundance distribution was significantly correlated with both the relative area of canopy gaps in the quadrats and the soil-topography gradient. Presumably, the critical factors involved in these two gradients are, respectively, light and ground water regimes. The influence of canopy gaps (i.e., light) was surprising and has not been documented previously for tropical deciduous dry forests.