Biomass,Carbon, and Nitrogen Pools in Mexican Tropical Dry Forest Landscapes

Tropical dry forest is the most widely distributed land-cover type in the tropics. As the rate of land-use/land-cover change from forest to pasture or agriculture accelerates worldwide, it is becoming increasingly important to quantify the ecosystem biomass and carbon (C) and nitrogen (N) pools of both intact forests and converted sites. In the central coastal region of México, we sampled total aboveground biomass (TAGB), and the N and C pools of two floodplain forests, three upland dry forests, and four pastures converted from dry forest. We also sampled belowground biomass and soil C and N pools in two sites of each land-cover type. The TAGB of floodplain forests was as high as 416 Mg ha^–1, whereas the TAGB of the dry forest ranged from 94 to 126 Mg ha^–1. The TAGB of pastures derived from dry forest ranged from 20 to 34 Mg ha^–1. Dead wood (standing and downed combined) comprised 27%–29% of the TABG of dry forest but only about 10% in floodplain forest. Root biomass averaged 32.0 Mg ha^–1 in floodplain forest, 17.1 Mg ha^–1 in dry forest, and 5.8 Mg ha^–1 in pasture. Although total root biomass was similar between sites within land-cover types, root distribution varied by depth and by size class. The highest proportion of root biomass occurred in the top 20 cm of soil in all sites. Total aboveground and root C pools, respectively, were 12 and 2.2 Mg ha^–1 in pasture and reached 180 and 12.9 Mg ha^–1 in floodplain forest. Total aboveground and root pools, respectively, were 149 and 47 kg ha^–1 in pasture and reached 2623 and 264 kg ha^–1 in floodplain forest. Soil organic C pools were greater in pastures than in dry forest, but soil N pools were similar when calculated for the same soil depths. Total ecosystem C pools were 306. The Mg ha^–1 in floodplain forest, 141 Mg ha^–1 in dry forest, and 124 Mg ha^–1 in pasture. Soil C comprised 37%–90% of the total ecosystem C, whereas soil N comprised 85%–98% of the total. The N pools lack of a consistent decrease in soil pools caused by land-use change suggests that C and N losses result from the burning of aboveground biomass. We estimate that in México, dry forest landscapes store approximately 2.3 Pg C, which is about equal to the C stored by the evergreen forests of that country (approximately 2.4 Pg C). Potential C emissions to the atmosphere from the burning of biomass in the dry tropical landscapes of México may amount to 708 Tg C, as compared with 569 Tg C from evergreen forests.     

Phenology of Tree Species in Bolivian Dry Forests

Phenological characteristics of 453 individuals representing 39 tree species were investigated in two dry forests of the Lomerío region, Department of Santa Cruz, Bolivia. The leaf, flower, and fruit production of canopy and sub–canopy forest tree species were recorded monthly over a two–year period. Most canopy species lost their leaves during the dry season, whereas nearly all sub–canopy species retained their leaves. Peak leaf fall for canopy trees coincided with the peak of the dry season in July and August. Flushing of new leaves was complete by November in the early rainy season. Flowering and fruiting were bimodal, with a major peak occurring at the end of the dry season (August–October) and a minor peak during the rainy season (January). Fruit development was sufficiently long in this forest that fruiting peaks actually tended to precede flowering peaks by one month. A scarcity of fruit was observed in May, corresponding to the end of the rainy season. With the exception of figs (Ficus), most species had fairly synchronous fruit production. Most canopy trees had small, wind dispersed seeds or fruits that matured during the latter part of the dry season, whereas many sub–canopy tree species produced larger animal– or gravity–dispersed fruits that matured during the peak of the rainy season. Most species produced fruit annually. Lomerio received less rainfall than other tropical dry forests in which phenological studies have been conducted, but rainfall can be plentiful during the dry season in association with the passage of Antarctic cold fronts. Still, phenological patterns in Bolivian dry forests appear to be similar to those of other Neotropical dry 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.     

Reproductive Traits and Phenology of Plants in Tropical Dry Evergreen Forest on the Coromandel Coast of India

Qualitative reproductive traits of 84 plant species belonging to 41 families were studied in tropical dry evergreen forest on the Coromandel coast of India. Majority of species had rotate type, white-coloured, scented flowers, rewarding nectar and pollen and pollinated chiefly by bees. An association between floral traits and pollination spectrum is evident. Bee pollination was prevalent in pollination systems. Among the fruit types, drupe and berry were common in black and red colour respectively, and dispersed by zoochorous mode. Seeds of brown- and green-coloured dry fruits, without any reward were disseminated by wind and explosion. The reproductive phenophase of trees and lianas occurred mostly during the dry period from January to June, which receives rainfall of less than 50 mm a month. However, shrubs showed a peak in flowering and fruiting in wet period. Detailed phenological observations of 22 woody species revealed a seasonal and unimodal pattern in flowering. Although some species were in flower round the year, flowering activity was skewed towards the dry season. The fruiting activity showed a bimodal pattern, one peak in dry season and another in wet season. Many species displayed a temporal aggregation in flowering and fruiting. The significant relation was obtained between reproductive traits and phenology of plants in the tropical dry evergreen forest.     

Microclimate Change and Effect on Fire Following Forest-Grass Conversion in Seasonally Dry Tropical Woodland1

We tested the hypothesis that, where fire has historically been infrequent, wooded areas that have been invaded by grasses and converted to grassland by fire are predisposed to future fire compared to adjacent areas that remain wooded; thus, an initial forest fire may promote future fires. We compared microclimate between a grass-dominated burned area and a nearby grass-invaded woodland that has not burned in recent history, both located in the submontane dry forest of Hawaii Volcanoes National Park. The results were used to parameterize BEHAVE, a fire behavior prediction model developed by the USDA Forest Service. The model's predictions include probability of ignition, intensity, rate of spread, and tree mortality. Contrary to expectations, daytime hourly mean temperatures were higher and relative humidity was lower in the woodland site. However, the differences in temperature and humidity were not great enough to affect spread rate or probability of ignition. Wind speeds were substantially greater in the grassland, and this was most important in driving differences in modeled fire spread. Given similar synoptic conditions, a fire started in the grassland can be expected to spread an order of magnitude faster than one started in the woodland.     

Litter Nutrient Dynamics During Succession in Dry Tropical Forests of the Yucatan: Regional and Seasonal Effects

Land-use change in the tropics is creating secondary forest at an unprecedented rate. In the tropical Americas, mature dry tropical forest is rapidly being converted to secondary forest during the fallow period of shifting cultivation. We investigated litter phosphorus (P) and nitrogen (N) dynamics in forests recovering from shifting cultivation of maize (corn) in three regions of the Southern Yucatan Peninsula, Mexico. Our goal was to understand how nutrient and water availability affect forest recovery following conversion of mature forest to agricultural land. To investigate such changes at a regional scale, newly fallen litter was collected monthly along a seasonal, a successional, and a precipitation gradient. Reflecting possible P limitation, litter P concentration declined with forest age, while litter N concentration did not differ between age classes. Average litter P concentration from the southern, wettest region was 0.87 mg/g, almost twice the litter P concentration in the drier central and northern regions (0.44 and 0.45 mg/g, respectively). Average N concentrations of litter from the three regions ranged from 1.1% to 1.2%, with no regional differences. However, minima in both P and N concentration from all regions were pronouncedly timed with peak litterfall, suggesting nutrient retranslocation during periods of water stress. Additionally, successional differences in litter P were clearest during wetter months. P nutrient-use efficiency was lowest in the southern region and highest in the central and northern study regions. N nutrient-use efficiency was up to 40 times lower than P nutrient-use efficiency and showed no regional differences. Overall, our results suggest that litter nutrient dynamics in secondary dry tropical forests of the Southern Yucatan are strongly influenced by water and nutrient availability, especially P, as well as land-use history.     

Seedling Establishment of a Canopy Tree Species in Malaysian Tropical Rain Forests

Abstract Unlike many other canopy tree species in tropical rain forests, Dryobalanops aromatica (Dipterocar-paceae, local name: Kapur) establishes monospecific dominant (monodominant) forests in Peninsular Malaysia. In natural conditions, monodominance of Kapur does not necessarily mean low species diversity of the Kapur forests. While the emergent canopy layer is occupied by Kapur, many other plant species, which are common to lowland dipterocarp primary forests in the same region, are found in lower canopy layer and understory.
To understand the ecological implications of the monodominance of Kapur, we monitored post dispersal survival and seedling establishment in a pure stand of Kapur in a plantation in Kepong, near Kuala Lumpur. Immediately after seed fall, seeds and cotyledon-stage seedlings suffered high predation by vertebrates such as rodents. The predation pressure was higher in a more general fruiting year (1991) than in a sporadic fruiting year (1992). In contrast to the high mortality of seeds and newly emerged seedlings, seedlings surviving to the six-leaf stage showed low mortality, which allowed the establishment of a sapling bank.
The occurrence of saplings of Kapur with a wide range of size classes in natural forests indicates that this species is more shade tolerant than other dipterocarp species such as the Shorea group and that it could well respond to enhanced light conditions caused by canopy opening. These characteristics may partly contribute to maintaining monodominance of Kapur.     

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.     

西双版纳热带季节雨林土壤氮含量对乔木树种多样性的影响?

大气氮沉降或人类活动导致生态系统氮输入增加,可能会提高土壤氮含量水平,促进优势种的生长和减少环境异质性,从而使物种共存的生态位减少,群落物种多样性降低。为研究土壤氮含量的增加对森林群落乔木树种多样性的影响,本研究在西双版纳热带季节雨林随机设置了14个1 ha的样方,对各样方土壤总氮( TN)含量、乔木树种丰富度以及西双版纳热带季节雨林20 ha动态监测样地中各样方乔木树种及建群种望天树( Parashorea chinensis)生物量进行了调查。结果表明：土壤氮含量与乔木树种丰富度具有显著负相关而与群落及建群种望天树生物量具有显著正相关。我们推测其机制可能是：土壤氮含量增加促进了建群种望天树等的生长及群落生物量的积累,减少树种共存的生态位,由于竞争排斥等原因而导致群落树种丰富度降低。因此,减少生态系统人为氮输入,对于保护西双版纳热带季节雨林乔木树种多样性具有重要意义。     

Species Richness,Spatial Variation,and Abundance of the Soil Seed Bank of a Secondary Tropical Rain Forest1

Despite the importance of the soil seed bank in tropical forest regeneration, little is known about spatial variability in species composition and abundance of seeds stored in the soil. To develop sampling methods for comparative studies, we examined species richness, spatial variation, and abundance of germinants from the soil seed bank in a 16 year old secondary, tropical wet forest at La Selva Biological Station, Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at the intersection points of a gridded 1 ha plot (10 × 10-m grid, 121 samples) and in a nested 100 m² subplot (2 × 2-m grid, 36 samples). The 1 ha plot had a density of 4535 seeds/m² with 34 species observed. Based on a series of 100 randomized species accumulation curves, a Michaelis-Menten fit predicted a mean species richness of 36.3 species; the number of observed species was close to the predicted asymptote. A nonparametric, first-order jackknife species richness estimator predicted a species richness of 37.0 species. Eighty-five and 95 percent of the observed species richness is contained, on average, within 41 and 74 pooled samples, respectively. Within the 100 m² nested subplot, a density of 5476 seeds/m² was observed, comprising 26 species with an estimated species richness (Michaelis-Menten fit) of 29.1 species. The jackknife species richness estimator predicted a species richness of 36.7 species. For species richness and abundance of both plots, spatial autocorrelation statistics (Moran's I) were not significantly different from zero at lag distances from 2 to 100 m, indicating a random distribution at these spatial scales. For this site, accurate estimates of species composition depend upon the number of samples collected as well as the spatial distribution of sampling effort. Many small samples distributed over a large area provide greater accuracy and precision for estimating species richness of the soil seed bank.     

Root Morphology and Anchorage of Six Native Tree Species from a Tropical Montane Forest and an Elfin Forest in Ecuador

Root architecture of tree species was investigated at two different altitudes in tropical forests in Ecuador. Increasing altitude was accompanied by higher wind speeds and more shallow soils, while slope angles of both sites were comparable (20–50°). Three tree species typical for the montane forest at 1900 m (Graffenrieda emarginata (Ruiz & Pav.) Triana (Melastomataceae), Clethra revoluta (Ruiz & Pav.) Spreng. (Clethraceae), Vismia tomentosa Ruiz & Pav. (Clusiaceae)) and for the elfin forest at 3000 m (Weinmannia loxensis Harling (Cunoniaceae), Clusia spec. (Clusiacaea) Styrax foveolaria Perkins (Styraceae)) were examined. At 1900 m, 92% of the trees grew upright, in comparison to 52% at 3000 m. At 3000 m, 48% of the trees were inclined, lying or even partly uprooted. At this altitude, all trees with tap roots or with shoots connected by coarse rhizomes, 83% of the trees with stilt roots, and 50% of the trees in which stems or roots were supported by other trees grew upright, suggesting that these characteristics were relevant for tree stability. Root system morphology differed markedly between altitudes. In contrast to 1900 m, where 20% of structural roots originated in the deeper mineral soil, root origin at 3000 m was restricted to the forest floor. The mean ratio of root cross sectional area to tree height decreased significantly from 6.1 × 10⁻³ m² m⁻¹ at 1900 m to 3.2 × 10⁻³ m² m⁻¹ at 3000 m. The extent of root asymmetry increased significantly from 0.29 at 1900 m to 0.62 at 3000 m. This was accompanied by a significantly lower number of dominant roots at 3000 m (2.3 compared to 3.8 at 1900 m). In conclusion, native tree species growing in tropical montane and elfin forests show a variety of root traits that improve tree stability. Root system asymmetry is less important for tree stability where anchorage is provided by a deep and solid root–soil plate. When deep rooting is impeded, root traits improving the horizontal extension of the root–soil plate are more pronounced or occur more frequently. Furthermore, mutual mechanical support of roots and stems of neighboring trees seems to be an appropriate mechanism to provide anchorage in soils with low bulk density and in environments with high wind speeds.     

香港次生林树种种子生理生态学特性的研究

１９８９～１９９１年在温室和野外测定了香港４０种次生林树木种类的种子寿命和发芽率．结果显示,在所测试的４０种天然林树种中,６０％的树种的种子寿命短于１年,仅有８个树种的种子寿命在１年以上．种子的寿命与种子在土壤中的发芽能力呈负相关的关系．种子发芽测定结果表明,大多数树种都可以在强光照条件（９５％的日光）下发芽,但３５％的测试树种不能在天然林下（０７％～２％的日光）发芽．最后从植物生理生态的角度,探讨种子特征与香港次生林区系发展及香港现有次生林的演替现状     

Predicting Species Response to Disturbance from Size Class Distributions of Adults and Saplings in a Jamaican Tropical Dry Forest

The diversity of tropical dry forests is poorly described and their regeneration ecology not well understood, however they are under severe threat of conversion and degradation. The Hellshire Hills constitute a dry limestone forest reserve on the south coast of Jamaica that is of high conservation value. In order to describe the structure and composition of this forest and assess the extent to which the population structures of its tree species do characterize their regeneration ecologies, pre-disturbance structure, floristics and seedling populations were compared with post-disturbance species responses in twelve 15 m × 15 m permanent sample plots which were laid out in a blocked design in April 1998, giving a total sample area of 0.27 ha. These plots were subjected to disturbance in April 1999 (cutting) with each of four blocks being assigned with two randomly allocated treatment plots (partially and clear cut) and one control plot (uncut). A total of 1278 trees (≥2 cm DBH) and 7863 seedlings and saplings (0–2 mm and 2–20 mm root collar diameter (RCD) respectively), comprising 60 and 52 species, respectively, were sampled in the plots prior to disturbance. The species-area curve for trees reached a maximum at 0.20 ha, and abundance was widely distributed amongst the species (26 had importance values greater than 1%); four species were notably codominant (with importance values between 7 and 8%). The forest stand structure had a reverse J-shaped curve for tree and for seedling/sapling size-class distributions, which indicated that the forest as a whole was probably regenerating adequately. From an analysis based on adult and sapling size-class distributions (SCDs), 21 species with 15 or more individuals were classified into 3 groups. Many of the species (15 of the 21), had flat adult SCDs that deviate from the whole-community reverse J-shaped SCD. However, sapling SCDs for 6 of the 15 species were strongly positive indicating the potential for their populations to be sustained by recruitment from the saplings present. No general association was found between these SCD species groupings and the actual ability of the species to recover from disturbance. Analysis of post-disturbance response revealed that for only 9 of the 21 species did adult SCDs provide adequate prediction, but for an additional 6 of the species information on sapling SCDs improved the accuracy of prediction if the ‘release’ of saplings or smaller individuals predominated recovery. However in this forest, recovery following disturbance which left stem and roots in place is predominantly by coppice regrowth, and there were no significant correlations found between adult SCDs and the species’ ability to coppice.     

Tree Regeneration and Species Diversity Following Conventional and Uniform Spacing Methods of Selective Cutting in a Subtropical Humid Forest Reserve1

There are a variety of ways to diminish the negative impacts of forest management for timber on biodiversity. A pilot project using a uniform spacing method of selective cutting was recently implemented in the 5340 ha Guarani Reserve, Misiones, Argentina, to design adequate management schemes for the subtropical forests of the region. Uniform spacing involves moderate timber harvesting and careful selection of remnant trees. In this article we compare tree regeneration in forests experimentally cut by diameter limit and uniform spacing methods. Seedlings of five size classes from 10 cm to 3 m in height were sampled using rectangular nested plots. Three years after cutting, an average of 54,330 and 22,270 seedling/ha (all height classes combined) of commercial and non-commercial species were found in the forest cut by uniform spacing and by minimum diameter, respectively. In an adjacent uncut forest there were 34,900 seedlings/ha, and in another forest cut by minimum diameters 30 years ago there were 50,000 seedlings/ha, The forest cut by uniform spacing had the highest number of commercial seedlings/ha, with three times as many as the forest cut by minimum diameter three years ago and twice as many as the forest treated by minimum diameters 30 years ago. The forest cut by uniform spacing and the forest treated by minimum diameters 30 years ago had the highest diversity of understory plants other than trees, as well as heterogeneous canopy cover, while the other two forests had a predominance of bamboo in the understory and more open canopy conditions. Although lack of site replication limits interpretation of the results, these early findings suggest that the uniform spacing method can be an ecologically sound forest management option for the region.     

Species Composition,Physiognomy and Plant Diversity of the Tropical Montane Evergreen Broad-leaved Forest in Southern Yunnan

Species composition, physiognomy and plant diversity of the less known tropical montane forests in southern Yunnan were studied based on the data from 15 sampling plots in three sites. These forests are mainly dominated by the families Theaceae, Fagaceae, Lauraceae and Euphorbiaceae in floristic composition, and dominated by evergreen phanerophytes with mesophyllous leaves. These forests are similar to lower montane rain forests in equatorial southeastern Asia in floristic composition and altitudinal distributions, but differ in physiognomy by having few epiphytes, but more lianas and more plants with compound leaves. These differences could be due to strongly seasonal climate and so-called mass elevation effect in southern Yunnan. They also differ from the tropical seasonal rain forests at lower altitudes in southern Yunnan by having conspicuously lower species richness, few epiphytes, fewer mega-mesophanerophytes, more abundant micro-nanophanerophytes and hemicryptophytes and more plants with microphyllous leaves. It is suggested that these forests could be termed tropical montane evergreen broad-leaved forests, and be a vegetation type from the northern margin of mainland southeastern Asia controlled by a strongly seasonal climate.     

The Role of Landscape Patterns of Habitat Types on Plant Species Diversity of a Tropical Forest in Mexico

The relationships among landscape characteristics and plant diversity in tropical forests may be used to predict biodiversity. To identify and characterize them, the number of species, as well as Shannon and Simpson diversity indices were calculated from 157 sampling quadrats (17,941 individuals sampled) while the vegetation classes were obtained from multi-spectral satellite image classification in four landscapes located in the southeast of Quintana Roo, Mexico. The mean number of species of trees, shrubs and vines as well as the mean value of the total number of species and the other two diversity indices were calculated for four vegetation classes in every one of the four landscapes. In addition, the relationships between landscape patterns metrics of patch types and diversity indices were explored. The multiple statistical analyses revealed significant predictor variables for the three diversity indices. Moreover, the shape, similarity and edge contrast metrics of patch types might serve as useful indicators for the number of species and the other two diversity variables at the landscape scale. Although the association between the three diversity indices and patch types metrics showed similar behavior, some differences were appreciated. The Shannon diversity index, with its greater sensitivity to rare species, should be considered as having a greater importance in interpretation analysis than Simpson index.