Phylogenetic structure of angiosperm communities during tropical forest succession

The phylogenetic structure of ecological communities can shed light on assembly processes, but the focus of phylogenetic structure research thus far has been on mature ecosystems. Here, I present the first investigation of phylogenetic community structure during succession. In a replicated chronosequence of 30 sites in northeastern Costa Rica, I found strong phylogenetic overdispersion at multiple scales: species present at local sites were a non-random assemblage, more distantly related than chance would predict. Phylogenetic overdispersion was evident when comparing the species present at each site with the regional species pool, the species pool found in each age category to the regional pool or the species present at each site to the pool of species found in sites of that age category. Comparing stem size classes within each age category, I found that during early succession, phylogenetic overdispersion is strongest in small stems. Overdispersion strengthens and spreads into larger size classes as succession proceeds, corroborating an existing model of forest succession. This study is the first evidence that succession leaves a distinct signature in the phylogenetic structure of communities.     

Plant functional traits under cattle grazing and fallow age scenarios in a tropical dry forest of Northwestern Mexico

Evaluating plant functional traits helps to understand how plants respond to changing environmental conditions and resource availability associated with disturbance events. Livestock production is one of the primary drivers of tropical forest loss and degradation. Livestock alter environmental conditions within the forest by grazing, trampling and nutrient inputs, which in turn can influence species composition and functional traits of species. Understanding how livestock influence functional traits along a successional gradient is poorly understood. Here, we studied the effect of cattle grazing and fallow age on plant functional traits and soil nutrients in secondary and old-growth tropical dry forests. We analyzed plant functional traits of the most important species in successional and old-growth forest communities in both cattle present and cattle excluded plots. Our results showed the effects of cattle grazing and fallow age on plant functional traits, with fallow age explaining more variation than cattle grazing. In early succession, functional traits were associated with water conservation (thicker leaves, lower specific leaf area), and in later successional they were linked with sunlight conservation (larger height, higher specific leaf area). The presence of large fruits and seeds in advanced successional sites suggests high resource availability, which may help plants to successfully reproduce. Moreover, under cattle grazing some functional traits are associated with herbivory defense (high foliar dry weight and thick leaves). Even though N and C increased as succession advanced, the sites with cattle grazing had higher NH4 and NO3 concentrations as a result of fecal deposition. Plant functional traits responded to fallow age than to cattle grazing. Our study showed that cattle exclusion, as a management and biodiversity conservation strategy, contributes positively to soil nutrition. Thus, fallow age and cattle exclusion facilitate soil recovery and allows establishing species with suitable functional attributes for overcoming environmental filters in abandoned cattle fields.     

Small-scale patterning in the recruitment of forest species during succession in tropical dry forest,Mozambique

Within a few years of field abandonment on the coastal dunes, a considerable number of species of trees and woody lianes of the climax evergreen forest enter the succession. At a relatively large-scale of analysis (a 100 m² plot), there is no evidence of soil development during the early succession, with no accumulation of soil organic matter nor nutrients being detected. At this scale there is, therefore, little support for the facilitation model of succession. However, when smaller scales of analysis are employed (e.g. a plot equivalent to the canopy area of a sapling) a very different picture emerges; most recruitment of forest woody species occurs beneath saplings found in early succession. Nutrient concentrations were higher in soils under sapling canopies than in soils beyond sapling canopies. With increasing sapling size, an increasing number of seedlings of forest woody species were found and there is progressive soil amelioration. Thus, during the early succession, there is considerable small-scale patterning in soil development and recruitment. This pattern is largely established through bird dispersal, with the intensity of recruitment being dependent on the distance to seed sources.     

Biomass and carbon accumulation in a fire chronosequence of a seasonally dry tropical forest

Seasonally dry tropical forests (SDTF) are a widely distributed vegetation type in the tropics, characterized by seasonal rainfall with several months of drought when they are subject to fire. This study is one of the first attempts to quantify above- and belowground biomass (AGB and BGB) and above- and belowground carbon (AGC and BGC) pools to calculate their recovery after fire, using a chronosequence approach (six forests that ranged form 1 to 29 years after fire and mature forest). We quantified AGB and AGC pools of trees, lianas, palms, and seedlings, and BGB and BGC pools (Oi, Oe, Oa soil horizons, and fine roots). Total AGC ranged from 0.05 to nearly 72 Mg C ha⁻¹, BGC from 21.6 to nearly 85 Mg C ha⁻¹, and total ecosystem carbon from 21.7 to 153.5 Mg C ha⁻¹; all these pools increased with forest age. Nearly 50% of the total ecosystem carbon was stored in the Oa horizon of mature forests, and up to 90% was stored in the Oa-horizon of early successional SDTF stands. The soils were shallow with a depth of <20 cm at the study site. To recover values similar to mature forests, BGC and BGB required <19 years with accumulation rates greater than 20 Mg C ha⁻¹ yr⁻¹, while AGB required 80 years with accumulation rates nearly 2.5 Mg C ha⁻¹ yr⁻¹. Total ecosystem biomass and carbon required 70 and 50 years, respectively, to recover values similar to mature forests. When belowground pools are not included in the calculation of total ecosystem biomass or carbon recovery, we estimated an overestimation of 10 and 30 years, respectively.     

The role of frugivorous bats in tropical forest succession

Discussion of successional change has traditionally focused on plants. The role of animals in producing and responding to successional change has received far less attention. Dispersal of plant propagules by animals is a fundamental part of successional change in the tropics. Here we review the role played by frugivorous bats in successional change in tropical forests. We explore the similarities and differences of this ecological service provided by New and Old World seed-dispersing bats and conclude with a discussion of their current economic and conservation implications. Our review suggests that frugivorous New World phyllostomid bats play a more important role in early plant succession than their Old World pteropodid counterparts. We propose that phyllostomid bats have shared a long evolutionary history with small-seeded early successional shrubs and treelets while pteropodid bats are principally dispersers of the seeds of later successional canopy fruits. When species of figs (Ficus) are involved in the early stages of primary succession (e.g. in the river meander system in Amazonia and on Krakatau, Indonesia), both groups of bats are important contributors of propagules. Because they disperse and sometimes pollinate canopy trees, pteropodid bats have a considerable impact on the economic value of Old World tropical forests; phyllostomid bats appear to make a more modest direct contribution to the economic value of New World tropical forests. Nonetheless, because they critically influence forest regeneration, phyllostomid bats make an important indirect contribution to the economic value of these forests. Overall, fruit-eating bats play important roles in forest regeneration throughout the tropics, making their conservation highly desirable.     

The role of remnant forest trees in tropical secondary succession

This paper emphasizes the contribution of remnant trees to the establishment of woody species during succession on abandoned fields and pastures in the Mexican rain forest area, Los Tuxtlas, Veracruz. Remnant trees, original large forest trees left in the clearings by traditional farmers will become natural perching sites for both passing and resident birds. Frugivorous birds drop or regurgitate seeds and fruits which fall under the canopies of remnant trees during their stay, thus contributing to an accumulation of species, which make these remnant trees into ‘regeneration nuclei’. The species transported into these sites belong chiefly to older stages of successional development and reach these otherwise isolated areas, counteracting the depauperization of tropical land, brought about by both intensive and extensive clearing. In a study of seven remnant trees, 29 woody species and two climbers were found, 86% of which are bird dispersed. The total number of species per tree varied from 6 to 15 and was higher under remnant trees with fruits attractive to birds. Floristic variations of the understorey as detected by detrended correspondence analysis was correlated with the relative amount of shade-tolerant primary and late secondary trees versus light dependent pioneers and early successional trees.     

中国西南地区热带森林演替序列碳动态

热带森林的破坏是全球性问题,我国西双版纳森林覆盖率受砍伐、火烧和短期耕种丢荒后,面积不断减少,取而代之的是大面积的不同演替状态的次生林。次生林演替过程中的碳储量和碳平衡的变化目前还鲜有研究,为了进一步揭示我国西南地区热带森林演替对于碳蓄积的影响,并制定更科学的热带森林经营管理措施,以结构复杂、生物多样和生物量巨大的热带森林为研究对象,并利用3个热带次生林的样地的实测数据,探讨了不同演替状态的热带次生林的碳储量变化,以及森林的净碳蓄积,死亡碳损失和更新碳增长等碳动态规律,分析表明:(1)在森林的演替过程中,森林的胸径分布频度从近正态分布逐渐向小径级的偏态分布发展,也就是随着演替的进展,小径级林木所占的比例越来越高。(2)热带次生林在森林固碳方面发挥着不可忽略的作用。(3)小的干扰,会波及森林的碳动态;大的干扰,如火灾和砍伐,将导致森林的次生演替,对森林的碳动态产生不可逆转的改变。(4)干旱事件是影响凋落物的季节和年间动态的原因,也是短时间尺度上影响碳平衡的一个重要因子。(5)不论原生林还是次生林,大树在生态系统碳动态方面皆扮演着重要的角色,因此本研究推荐注重大树的研究。     

Root biomass of a dry deciduous tropical forest in Mexico

The deciduous tropical dry forest at Chamela (Jalisco, Mexico) occurs in a seasonal climate with eight rainless (November through June) and four wet months (700 mm annual precipitation). The forest reaches a mean height of 10 m. Tree density in the research area was 4700 trees per ha with a basal area at breast height of 23 m² per ha. The above-and below-ground biomass of trees, shrubs, and lianas was 73.6 Mg ha^–1 and 31 Mg ha^–1, respectively. A root:shoot biomass ratio of 0.42 was calculated. Nearly two thirds of all roots occur in the 0–20 cm soil layer and 29% of all roots have a diameter of less than 5 mm.     

Secondary succession in abandoned fields of dry tropical Northern Cameroon

Abstract. This study evaluates the processes of recolonization of abandoned fields by native vegetation under conditions of intensive human activity (fire, intensive grazing, firewood cutting) in a semi-arid tropical region savanna of northern Cameroon. Secondary plant succession was studied in two series of formerly cultivated fields 1–35 yr after the beginning of the fallow period. Floristic changes and the dynamics of woody plant populations were compared between areas with vertisols (clay texture) and sandy soils, as a function of length of fallows. Vegetation changed continuously during the 35 years following field abandonment. However, a very abrupt break occurred between 6 and 10 yr, due to increasingly intense human pressure during this period. Up to that point, ecological models and mechanisms of succession presented in the literature are more or less confirmed by our results. Usually, secondary succession is blocked at a stage of wooded grassland as a result of human activities.     

缙云山森林群落次生演替中土壤理化性质的动态变化

选择典型、有代表性的不同演替阶段群落,进行了植被调查和土壤分析.结果表明,土壤理化性质在演替方向和土壤剖面上表现出较强的规律.土壤有机质随植被从低级向高级演进逐渐积累,分别是19.5(X₁)、37.0(X²)、50.1(X₃)和71.6 g·kg^-1(X₄);土壤全N、碱解氮和速效钾等也呈上升趋势;土壤pH和盐基饱和度降低,阳离子交换量增加.在土壤剖面上,有机质、全N等指标表现出A＞B＞C层的趋势.灰色关联度分析表明,随着演替的进行,土壤肥力提高,物种丰富度和郁闭度也相应增加.不同群落土壤理化性质在不同季节有显著差异,但这种差异并不影响植物与土壤在大时间尺度下的演变方向.土壤理化性质的动态变化与植物演替相适应.     

热带人工林土壤动物群落的次生演替和发展过程探讨

在广东小良地区的桉林迹地上,对不同林龄的阔叶混交林土壤动物的组成进行比较,探讨群落随林龄的增加而变化的过程。以多样性为基础,可将这里土壤动物群落的演替和发展分成3个阶段,即增长期、过渡期和稳定期。在林龄20多年后,群落发展可进入最后阶段。不同类群的演变程序:(1)湿生动物个体数逐渐增加,直至平稳;(2)昆虫中不稳定类群逐渐减少;(3)白蚁种群在增长期前段迅速增加,随后即在较低水平上受到控制;(4)蚯蚓在增长期后段才出现,并迅速成为优势类群。过渡期及其以前的蚯蚓种类主要为寒(虫宪)蚓属(Ocnerodrilus),稳定后为环毛蚓属(Pheretima)。     

Effects of herbivory and the season of disturbance on algal succession in a tropical intertidal shore,Phuket, Thailand

The effects of herbivory and the season of disturbance on species composition and algal succession were experimentally tested at a tropical intertidal shore, Phuket Island, Thailand. Dead coral patches were cleared of all organisms during both the dry and rainy seasons in order to study the effects of season on algal succession and cages were set up to exclude fish herbivory. Algal succession in this intertidal habitat showed a simple pattern and took a year from the early Ulva paradoxa C. Agardh stage to the late Polysiphonia sphaerocarpa Børgesen stage. The abundance of algae during succession was under the influence of seasonal change. U. paradoxa reproduced and recruited throughout the year. Caging effects did not apparently influence algal abundance, perhaps because resident herbivorous damselfishes excluded other herbivores from their territories and maintained their algal “farms”. Unexpectedly, the percent cover of Ulva in the caged plots was lower than in uncaged plots. This pattern may indicate that caging excluded damselfishes only, but allowed small herbivores that consumed substantial amounts of soft filamentous algae in the cages.     

Temporal variation in the influence of forest succession on caterpillar communities: A long‐term study in a tropical dry forest

Forest succession can influence herbivore communities through changes in host availability, plant quality, microclimate, canopy structure complexity and predator abundance. It is not well known, however, if such influence is constant across years. Caterpillars have been reported to be particularly susceptible to changes in plant community composition across forest succession, as most species are specialists and rely on the presence of their hosts. Nevertheless, in the case of tropical dry forests, plant species have less defined successional boundaries than tropical wet forests, and hence herbivore communities should be able to persist across different successional stages. To test this prediction, caterpillar communities were surveyed during eight consecutive years in a tropical dry forest in four replicated successional stages in Chamela, Jalisco and Mexico. Lepidopteran species richness and diversity were equivalent in mature forests and early successional stages, but a distinctive caterpillar community was found for the recently abandoned pastures. Species composition tended to converge among all four successional stages during the span of eight years. Overall, our results highlight the importance of both primary and secondary forest for the conservation of caterpillar biodiversity at a landscape level. We also highlight the relevance of long‐term studies when assessing the influence of forest succession to account for across year variation in species interactions and climatic factors. Abstract in French is available with online material.     

Impact of logging on the richness and diversity of forest butterflies in a tropical dry forest in Thailand

Studies on the impact of logging on tropical forest butterflies have been almost exclusively conducted in moist forest habitats. This study considers the impacts of small-scale logging on butterfly communities at three sites of varying disturbance intensity in a tropical dry forest in western Thailand. Butterfly species richness was similar at all sites, but the abundance of butterflies and diversity of the butterfly community decreased with increased logging disturbance. The recorded decrease in diversity at the relatively large sampling scale used lends further support to the hypothesis that disturbance effects are scale dependent. Species abundance data for butterflies fitted a log-normal distribution at all sites, but also a log-series distribution at the two disturbed sites. These analyses suggest a more complex butterfly community at the undisturbed site, but also that log-series and log-normal distributions may not to be sufficiently sensitive to be useful indicators of community changes following logging. Community ordination separates both the butterfly species and transect samples into three distinct regions corresponding to the three study locations. Ordination axes are correlated with tree density, understorey cover and understorey plant richness. Species with the smallest geographic ranges tend to be the least abundant and occurred most frequently in the undisturbed site. The observed diverging responses to disturbance among butterfly families diminishes the value of butterfly communities as biodiversity indicators, and forest managers should perhaps focus on restricted range species or of groups of recognized sensitive species for this purpose.     

Flower biology of the cactus Coryphantha elephantidens in the tropical dry forest of Central Mexico

Floral traits and sexual systems in angiosperms are strategies that enhance outcrossing within hermaphrodite flowers and among individuals in a population. Sexual systems with unisexual flowers have also evolved among angiosperms, resulting in sex specialization. Furthermore, the interaction of floral traits and floral visitors determines successful plant reproduction. Globose cacti are bee pollinated, and variation in the diversity of their pollinator assemblages is strongly associated with floral phenotype. Our objective was to describe the floral biology of the cactus Coryphantha elephantidens and to determine its relationship with pollinators. Floral traits were studied by direct observations in live and fixed flowers. The breeding system was determined using two estimators based on floral morphology: pollen grains to ovules per flower (P/O) ratio and outcrossing index. Pollination treatments were conducted to determine the mating system. Floral visitors were recorded using direct observation. Flowers of C. elephantidens are variable in color, protandric, herkogamous and nectarless. Estimators of the breeding system indicated xenogamy, which is consistent with the obligate outcrossing revealed by the pollination experiment. Thirty-seven percent of the plants have female flowers that do not produce pollen, making this population functionally gynodioecious. Both fruit and seed set were high compared to other globose cacti. Pollinators included eight species of native bees, a more diverse pollinator assemblage than other globose cacti. Given the high pressure on pollen due to functional gynodioecy, nectarless flowers, an outcrossing mating system, and the necessity of pollinators to set seeds, we concluded that native bees are highly efficient pollinators that play a crucial role in the sexual reproduction of C. elephantidens.     

Validating distribution models for twelve endemic bird species of tropical dry forest in western Mexico

Considering the high biodiversity and conservation concerns of the tropical dry forest, this study aim is to predict and evaluate the potential and current distributions of twelve species of endemic birds which distribute along the western slope of Mexico. The main goal is to evaluate altogether different methods for predicting actual species distribution models (ADMs) of the twelve species including the identification of key environmental potential limiting factors. ADMs for twelve endemic Mexican birds were generated and validated by means of applying: (1) three widely used species niche modeling approaches (ENFA, Garp, and Maxent); (2) two thresholding methods, based on ROC curves and Kappa Index, for transforming continuous models to presence/absence (binary) models; (3) documented habitat–species associations for reducing species potential distribution models (PDMs); and (4) field occurrence data for validating final ADMs. Binary PDMs' predicted areas seemed overestimated, while ADMs looked drastically reduced and fragmented because of the approach taken for eliminating those predicted areas which were documented as unsuitable habitat types for individual species. Results indicated that both thresholding methods generated similar threshold values for species modeled by each of the three species distribution modeling algorithms (SDMAs). A Wilcoxon signed‐rank test, however, showed that Kappa values were generally higher than ROC curve for species modeled by ENFA and Maxent, while for Garp models there were no significant differences. Prediction success (e.g., true presences percentage) obtained from field occurrence data revealed a range of 50%–82% among the 12 species. The three modeling approaches applied enabled to test the application of two thresholding methods for transforming continuous to binary (presence/absence) models. The use of documented habitat preferences resulted in drastic reductions and fragmentation of PDMs. However, ADMs predictive success rate, tested using field species occurrence data, varied between 50 and 82%.     

Root system analysis of seedlings of seven tree species from a tropical dry forest in Mexico

Summary Root attributes of tree seedlings of seven species from the tropical deciduous forest along the Pacific Coast of Mexico are described using morphometirc root system analysis. Mean relative growth rate, root/shoot ratios, specific root length, root density, mean number of roots tips and root length/leaf area ratio were determined in seedlings grown for 35 days inside growth chambers. All the species had low relative growth rates, low root/shoot ratios and low root densities (<0.5 cm/cm³). The species associated with disturbed habitats, in contrast to the species characteristic of undisturbed areas, presented small seeds, a dichotomous root branching pattern and large specific root length. No relationship was found between seed size and mean relative growth rate among the species studied.     

Liana community and functional trait analysis in tropical dry evergreen forest of India

Aim Lianas are abundant and diverse throughout the world and constitute an important structural and functional component of tropical forests. This study aims to investigate liana diversity, abundance and their functional traits in Indian tropical dry evergreen forest (TDEF).Methods A total of ten 1-ha plots, one each in 10 Indian TDEF sites were demarcated. Each 1-ha plot was divided into one-hundred 10- × 10-m quadrats to facilitate woody species inventory. All lianas ≥1cm diameter measured at 130cm from the rooting point and all trees ≥10-cm girth at breast height (gbh) were recorded from the study sites to analyze the patterns of liana diversity and abundance and also to compare the contribution of lianas to the total woody species richness, density and basal area. Liana variables across the study sites were compared using one-way analysis of variance. The qualitative functional traits of inventoried lianas and trees were assessed on the field and referring to pertinent field manuals.Important findings A total of 9237 liana individuals (ranged from 408–1658 individuals ha^-1) representing 52 species, 45 genera and 28 families were encountered from the 10 study sites. Liana species richness ranged from 11–31 species ha^-1 in 10 sites, which averaged 23.4 (±5.7) species ha^-1. The total basal area of lianas in the study sites was 7. 3 m 2 (0.20–1.76 m 2 ha^-1). There was a significant variation in liana species richness, density and basal area across the studied sites. On the whole, lianas contributed 52%, 49.3% and 4.1% to the total woody species (lianas and trees) richness, density and basal area, respectively. Liana trait analysis revealed the majority (50%) of lianas belonged to brevi-deciduous type. Stem twining was the chief climbing mechanism, exhibited by 21 species (52.6% of total abundance). More than half of the liana species (34 species; 6925 individuals) had microphyllous leaves. Fleshy-fruited lianas mostly bearing berries and drupes constituted the major fruit type in the studied sites. Zoochory was the predominant dispersal mode observed in 63.4% of species. Considering the ecological and functional role of lianas in Indian TDEF, the need for conservation is emphasized.     

Thermal ecophysiology of a native and an invasive gecko species in a tropical dry forest of Mexico

For ectotherms, thermal physiology plays a fundamental role in the establishment and success of invasive species in novel areas and, ultimately, in their ecological interactions with native species. Invasive species are assumed to have a greater ability to exploit the thermal environment, higher acclimation capacities, a wider thermal tolerance range, and better relative performance under a range of thermal conditions. Here we compare the thermal ecophysiology of two species that occur in sympatry in a tropical dry forest of the Pacific coast of Mexico, the microendemic species Benedetti's Leaf-toed Gecko (Phyllodactylus benedettii) and the invasive Common House Gecko (Hemidactylus frenatus). We characterized their patterns of thermoregulation, thermoregulatory efficiency, thermal tolerances, and thermal sensitivity of locomotor performance. In addition, we included morphological variables and an index of body condition to evaluate their effects on the thermal sensitivity of locomotor performance in these species. Although the two species had similar selected temperatures and thermal tolerances, they contrasted in their thermoregulatory strategies and thermal sensitivity of locomotor performance. Hemidactylus frenatus had a higher performance than the native species, P. benedettii, which would represent an ecological advantage for the former species. Nevertheless, we suggest that given the spatial and temporal limitations in habitat use of the two species, the probability of agonistic interactions between them is reduced. We recommend exploring additional biotic attributes, such as competition, behavior and niche overlap in order assess the role of alternative factors favoring the success of invasive species.