Spatial and temporal variation in the biomass and nutrient status of epilithic algae in Lake Erken, Sweden

1. The aim of this study was to estimate patchiness in biomass and in the internal nutrient status of benthic algae on hard substrata (epilithon) in Lake Erken, Sweden, over different levels of distance, depth and time. Knowledge of the sources and scale of patchiness should enable more precise estimation of epilithic biomass and nutrient status for the entire lake. We focused on the horizontal scale, about which little is known. 2. We sampled epilithon by SCUBA diving and used a hierarchical sampling design with different horizontal scales (cm, dm, 10 m, km) which were nested in two temporal scales (within and between seasons). We also compared two successive years and three sampling depths (0, 1 and 4 m). Biomass was measured as particulate carbon and chlorophyll a (Chl a) and internal nutrient status as carbon : nitrogen : phosphorus (C : N : P) ratios and as specific alkaline phosphatase activity (APA). 3. Horizontal variation accounted for 60–80 and 7–70% of the total variation in biomass and in nutrient status, respectively, at all depths and during both years. Both small and large scales accounted for significant variation. We also found variation with time and depth. Biomass increased in autumn after a summer minimum, and the within‐season variation was very high. The lowest biomass was found at 0 m depth. Both N and P limitation occurred, being higher in 1996 than in 1997 and decreased with depth. 4. As a consequence, any sampling design must address variation with distance, depth and time when estimating biomass or nutrient limitation of benthic algae for an entire lake. Based on this analysis, we calculated an optimal sampling design for detecting change in the epilithic biomass of Lake Erken between different sampling days. It is important to repeat the sampling as often as possible, but also the large scales (10 m and km) and the dm scale should be replicated. Using our calculations as an example, and after a pilot study, an optimal sampling design can be computed for various objectives and for any lake. 5. Short‐term impact of the wind, light and nutrient limitation, and grazing, might be important in regulating the biomass and nutrient status of epilithic algae in Lake Erken. Patchiness in the nutrient status of algae was not coupled to the patchiness of biomass, indicating that internal nutrients and biomass were regulated by different factors.     

Phytoplankton succession in Lake Valencia,Venezuela

Phytoplankton counts and supporting physical and chemical data were taken on Lake Valencia, Venezuela, over a five-year interval. The data are used to test the validity of a successional paradigm for class-level taxa. According to the paradigm, formulated from previous studies of Lake Lanao, Philippines, and from data on temperate lakes, the order of taxa from early to late succession is: diatoms, chlorophytes, blue-green algae, dinoflagellates. A successional episode is considered to begin when stability of a water column is restored after deep mixing. As the episode progresses, there is a steady decrease in concentration of the limiting macronutrient (in this case, N). In a test of the validity of the paradigm for Lake Valencia, dates of exceptional population increase or decrease were obtained for each taxon. Since nitrate concentration declines steadily as succession progresses, the entry of a given taxon into the successional sequence is indicated quantitatively by the mean nitrate concentration on dates of exceptional increase in population density, and exit from the successional sequence is indicated by mean nitrate concentration on dates of exceptional population declines. The successional position of each major taxon, bounded by its entry and exit in the sequence, can be mapped on the complete spectrum of nitrate concentrations observed in the lake. For Lake Valencia, the nitrate mapping procedure agrees exactly with the predictions based on the successional paradigm. Conformance of Lake Valencia phytoplankton with predictions made a priori suggests that there is a generalized pattern in the phytoplankton succession of the mixed layers of temperate and tropical lakes.     

Phytoplankton succession and diversity in a warm monomictic,relatively shallow lake: Lake Volvi,Macedonia, Greece

In Lake Volvi, phytoplankton dominance was examined in relation to the main phases of the fluctuating physical state of the water column and nutrient levels. Four stages of algal succession were identified. The successional pattern was, in general, similar from year to year. External disturbances such as increased wind mixing and high floods had the effect of setting the succession back to an earlier stage.Nanoplanktic diatoms and flagellates dominated during the first stages of succession. The succession was running from r-selected species in early stages to K-strategists in summer and towards a mixed community in the terminal stage.Diversity was expressed by the indices of Gleason and Shannon (based on physical units and biomass). Interpretation of species diversity took into account the differential variations of its two components, the number of species and the evenness of their distribution. Diversity was more affected by evenness than by number of species.Diversity increased in late stages of succession corresponding to the complexity of the community (high number of species, high evenness). Physical disturbances influenced positively the diversity. The maximal diversity appeared in the transition periods between the compositional changes generated by disturbances and the true successional development. Disturbances may generate a more uniform distribution of diversity throughout the whole water column.The role of physical disturbances in increasing the phytoplankton diversity may be demonstrated from comparisons of diversity — evenness — species number of the same stages under different meteorological conditions.     

The relationship between tree biodiversity and biomass dynamics changes with tropical forest succession

Theory predicts shifts in the magnitude and direction of biodiversity effects on ecosystem function (BEF) over succession, but this theory remains largely untested. We studied the relationship between aboveground tree biomass dynamics (Δbiomass) and multiple dimensions of biodiversity over 8–16 years in eight successional rainforests. We tested whether successional changes in diversity–Δbiomass correlations reflect predictions of niche theories. Diversity–Δbiomass correlations were positive early but weak later in succession, suggesting saturation of niche space with increasing diversity. Early in succession, phylogenetic diversity and functional diversity in two leaf traits exhibited the strongest positive correlations with Δbiomass, indicating complementarity or positive selection effects. In mid‐successional stands, high biodiversity was associated with greater mortality‐driven biomass loss, i.e. negative selection effects, suggesting successional niche trade‐offs and loss of fast‐growing pioneer species. Our results demonstrate that BEF relationships are dynamic across succession, thus successional context is essential to understanding BEF in a given system.     

Seasonal Dynamics of Periphyton in a Large Tropical Lake

Tropical aquatic systems are generally assumed to have little seasonality in productivity patterns. However, this study indicated that there was substantial seasonal variation in epilithic productivity and biomass in tropical Lake Tanganyika, due primarily to seasonal patterns in lake hydrodynamics that influence nutrient availability. Although they support much of the lake’s biological diversity, epilithic algae made a minor contribution to the total energy budget in Lake Tanganyika. A comparison among large, oligotrophic lakes revealed no significant latitudinal trends in periphyton productivity or biomass. However, Lake Tanganyika has relatively low benthic algal biomass and is therefore more efficient at photosynthesis than the temperate lakes. The influence of wave action and consumer density and diversity may be important in moderating productivity of the epilithic community.     

Microbial community composition but not diversity changes along succession in arctic sand dunes

The generality of increasing diversity of fungi and bacteria across arctic sand dune succession was tested. Microbial communities were examined by high‐throughput sequencing of 16S rRNA genes (bacteria) and internal transcribed spacer (ITS) regions (fungi). We studied four microbial compartments (inside leaf, inside root, rhizosphere and bulk soil) and characterized microbes associated with a single plant species (Deschampsia flexuosa) across two sand dune successional stages (early and late). Bacterial richness increased across succession in bulk soil and leaf endosphere. In contrast, soil fungal richness remained constant while root endosphere fungal richness increased across succession. There was, however, no significant difference in Shannon diversity indices between early and late successional stage in any compartment. There was a significant difference in the composition of microbial communities between early and late successional stage in all compartments, although the major microbial OTUs were shared between early and late successional stage. Co‐occurrence network analysis revealed successional stage‐specific microbial groups. There were more co‐occurring modules in early successional stage than in late stage. Altogether, these results emphasize that succession strongly affects distribution of microbial species, but not microbial diversity in arctic sand dune ecosystem and that fungi and bacteria may not follow the same successional trajectories.     

Multifractal spatial patterns and diversity in an ecological succession

We analyzed the relationship between biodiversity and spatial biomass heterogeneity along an ecological succession developed in the laboratory. Periphyton (attached microalgae) biomass spatial patterns at several successional stages were obtained using digital image analysis and at the same time we estimated the species composition and abundance. We show that the spatial pattern was self-similar and as the community developed in an homogeneous environment the pattern is self-organized. To characterize it we estimated the multifractal spectrum of generalized dimensions D(q). Using D(q) we analyze the existence of cycles of heterogeneity during succession and the use of the information dimension D(1) as an index of successional stage. We did not find cycles but the values of D(1) showed an increasing trend as the succession developed and the biomass was higher. D(1) was also negatively correlated with Shannon's diversity. Several studies have found this relationship in different ecosystems but here we prove that the community self-organizes and generates its own spatial heterogeneity influencing diversity. If this is confirmed with more experimental and theoretical evidence D(1) could be used as an index, easily calculated from remote sensing data, to detect high or low diversity areas.     

Microbial succession of nitrate-reducing bacteria in the rhizosphere of Poa alpina across a glacier foreland in the Central Alps

Changes in community structure and activity of the dissimilatory nitrate-reducing community were investigated across a glacier foreland in the Central Alps to gain insight into the successional pattern of this functional group and the driving environmental factors. Bulk soil and rhizosphere soil of Poa alpina was sampled in five replicates in August during the flowering stage and in September after the first snowfalls along a gradient from 25 to 129 years after deglaciation and at a reference site outside the glacier foreland (>2000 years deglaciated). In a laboratory-based assay, nitrate reductase activity was determined colorimetrically after 24 h of anaerobic incubation. In selected rhizosphere soil samples, the community structure of nitrate-reducing microorganisms was analysed by restriction fragment length polymorphism (RFLP) analysis using degenerate primers for the narG gene encoding the active site of the membrane-bound nitrate reductase. Clone libraries of the early (25 years) and late (129 years) succession were constructed and representative clones sequenced. The activity of the nitrate-reducing community increased significantly with age mainly due to higher carbon and nitrate availability in the late succession. The community structure, however, only showed a small shift over the 100 years of soil formation with pH explaining a major part (19%) of the observed variance. Clone library analysis of the early and late succession pointed to a trend of declining diversity with progressing age. Presumably, the pressure of competition on the nitrate reducers was relatively low in the early successional stage due to minor densities of microorganisms compared with the late stage; hence, a higher diversity could persist in this sparse environment. These results suggest that the nitrate reductase activity is regulated by environmental factors other than those shaping the genetic structure of the nitrate-reducing community.     

镜泊湖岩溶台地不同植被类型土壤微生物群落特征

为了探讨不同演替阶段植被类型土壤微生物群落特征,分别选取镜泊湖岩溶台地草本、矮灌木、高灌木、小乔木与灌木混生(简称混生)群落、落叶阔叶林及针阔混交林6种典型植被类型,进行植物群落调查和对土壤微生物生物量、群落结构和多样性指标、土壤物理化学性质的测定。结果表明：从土壤微生物量、土壤微生物群落组成、土壤微生物代谢动力学过程和代谢功能多样性的角度来看,各种植被类型土壤微生物群落具有明显的差异。演替前期的草本群落土壤微生物量碳氮、细菌生物量、真菌生物量,代谢活性及丰富度指数均最低,但Shannon-Wiener多样性指数和均匀度指数显著(P<0.05)高于其他植被类型。矮灌木土壤微生物群落组成显著受植被类型的影响。高灌木群落和混生(小乔木与灌木混生)群落具有极强的相似性, 但在碳源利用类型上两者表现出一定的差异。落叶阔叶林代谢活性最高,碳源利用能力最强,能利用BIOLOG微孔板中的所有31种碳源,这与其具有较高的微生物量碳氮和细菌生物量一致,其代谢功能丰富度最高。演替后期的针阔混交林下的土壤pH最低,真菌比例升高,在碳源丰富的条件下具有极强的竞争优势(仅次于落叶阔叶林),但在碳源贫瘠的条件下其利用碳源能力较弱(仅高于草本)。植被可能主要通过土壤全磷和有机质影响土壤微生物代谢功能多样性。     

Epilithic chlorophyll a,photosynthesis, and respiration in control and fertilized reaches of a tundra stream

Photosynthesis and respiration by the epilithic community on cobble in an arctic tundra stream, were estimated from oxygen production and consumption in short-term (4–12 h), light and dark, chamber incubations. Chlorophyll a was estimated at the end of each incubation by quantitatively removing the epilithon from the cobble. Fertilization of the river with phosphate alone moderately increased epilithic chlorophyll a, photosynthesis, and respiration. Fertilization with ammonium sulfate and phosphate, together, greatly increased each of these variables. Generally, under both control and fertilized conditions, epilithic chlorophyll a concentrations (mg m⁻²), photosynthesis, and respiration (mg O₂ m⁻², h⁻¹) were higher in pools than in riffles. Under all conditions, the P/R ratio was consistent at ∼ 1.8 to 2.0. The vigor of epilithic algae in riffles, estimated from assimilation coefficients (mg O₂ [mg Chl a]⁻¹ h⁻¹) was greater than the vigor of epilithic algae in pools. However, due to the greater accumulation of epilithic chlorophyll a in pools, total production (and respiration) in pools exceeded that in riffles. The epilithic community removed both ammonium and nitrate from water in chambers. Epilithic material, scoured by high discharge in response to storm events and suspended in the water column, removed ammonium and may have increased nitrate concentrations in bulk river water. However, these changes were small compared to the changes exerted by attached epilithon.     

Effects of earthworms and arbuscular mycorrhizal fungi depend on the successional stage of a grassland plant community

Earthworms and arbuscular mycorrhizal fungi (AMF) have profound impacts on plant performance. However, it is largely unknown if and how earthworms and AMF may affect plant succession. We planted mesocosms with an early-mid successional and a mid-late successional grassland plant community and added endogeic earthworms and commercial AMF in a full-factorial way to natural background soil. Earthworms had a positive effect on the total root and shoot biomass of both plant communities, with the effect on the shoots being slightly enhanced by co-inoculation with AMF. Surprisingly, the earthworm effect on the mid successional plant species depended on the successional stage of the plant community. Earthworms had a positive effect on the mid successional plant species when they were growing in the mid-late successional plant community, but no effect when the same plant species were growing in the early-mid successional plant community. Addition of AMF alone tended to reduce the shoot biomass of the early successional plant species, while the addition of earthworms in the presence or absence of AMF increased their shoot biomass. We conclude that the impacts of earthworms on plant species may depend on the successional stage of the plant community, while the effect of AMF addition depends on the successional stage of the plant community and may be changed by the presence of earthworms. Earthworms and AMF addition affect plants and plant communities of different successional stages differently with potential effects on plant succession.     

Environmental heterogeneity and spatiotemporal variability in plant defense traits

At the scale of the local plant community, we know very little about how spatial and temporal environmental heterogeneity affects the diversity in types and levels of plant defenses. If environmental heterogeneity is an important mechanism influencing plant defense traits, then defense expression should co‐vary spatially with environmental conditions and change as succession progresses. In this study, we examined how spatial heterogeneity and succession influence putative resistance and tolerance trait levels in late goldenrod Solidago altissima. We quantified the spatial distributions of herbivore damage and traits associated with resistance (leaf toughness, phenolics), tolerance (specific leaf area, relative growth rate, leaf addition rate and leaf senescence rate), and fitness (height, diameter, inflorescence biomass) of goldenrods within replicate early‐ and late‐successional fields. Also, we characterized the local neighborhood (stem density, canopy cover, ground vegetative cover) and edaphic conditions (soil moisture, pH, N) surrounding each target ramet, and determined relationships between these environmental variables and goldenrod trait levels. The distribution of traits within fields was strongly non‐random, and defense‐trait levels were more strongly spatially structured (i.e. autocorrelated) in late‐ than in early‐successional fields. Also, defense traits were most strongly correlated with aspects of the local plant neighborhood, and these relationships differed in important ways between successional stages. In late‐successional fields, tolerance trait specific leaf area was positively correlated with canopy cover and negatively correlated with stem density. In early‐successional fields, the relationship between ground vegetative cover and resistance (i.e. 1 – damage) was significantly stronger than in late‐successional fields. A novel insight from this study is the possibility that changes in the biotic environment during succession may shift the expression of defense from a resistance to a tolerance strategy in our system. This study highlights the context dependence of plant defense trait levels, which may promote their spatial and temporal variability in heterogeneous landscapes.     

Variations and Trade‐offs in Functional Traits of Tree Seedlings during Secondary Succession in a Tropical Lowland Rain Forest

The seedling stage is generally the most important bottleneck for the successful regeneration of trees in forests. The traits of seedlings, particularly biomass allocation and root traits, are more easily quantified than the traits of adults. In this study, we tested the hypothesis that seedling traits vary and trade‐off tracking the changing environment during secondary succession. We measured the major morphological traits of 27 dominant species and the major environmental factors in a chronosequence (30‐yr‐old fallow, 60‐yr‐old fallow, and old growth forest) after shifting cultivation in a tropical lowland rain forest on Hainan Island, China. The 30‐yr‐old fallow had higher light and nutrient availability, and the older forests had higher soil water content. Redundancy analysis based on species abundance and environmental factors revealed groups of seedlings that dominate in different stages of succession. Seedlings in different stages of succession had different strategies of biomass allocation for harvesting resources that varied in availability. Species characteristic of younger forest had higher allocation to roots and higher specific leaf area, while species characteristic of older forest had higher allocation to leaves. Our study suggests that the variations and trade‐offs in the major functional traits of tree seedlings among successional classes may reflect changes in environmental conditions during succession.     

Plant diversity patterns in subtropical evergreen broad-leaved forests of Yunnan and Taiwan

The subtropical evergreen broad-leaved forests of Yunnan and Taiwan were compared along environmental and successional gradients with the aim of identifying important taxon and species diversity as well as the drivers of mountain biodiversity patterns. A detrended correspondence analysis of an exhaustive set of data collected from 105 and 223 plots for Yunnan and Taiwan, respectively, was applied to classify natural mature forest types. Additional data from 72 and 68 plots for Yunnan and Taiwan, respectively, were used for analyses of secondary succession. The floristic richness and diversity index were calculated for each type of forest. In Yunnan, the monsoon forests in mesic-humid sites had more taxa and tended to show higher species diversity than the other two forest types. In Taiwan, species diversity values were significantly higher in the Machilus–Castanopsis zone in the middle altitudes (500–1500 m) than for the other three forest zones. For both Yunnan and Taiwan, the forests at the middle successional stage showed significantly higher species diversity than those at the early successional stage. Differences in diversity between the middle and late stages were not significant. These findings highlight the high species diversity of the natural mature evergreen broad-leaved forests of both Yunnan and Taiwan. In the secondary forests, as succession proceeds, species diversity comes to resemble that of the natural mature forests. In both ecosystems, the drivers of species diversity patterns are moisture, altitude, and succession/disturbance.     

冰川消退带微生物群落演替及生物地球化学循环

冰川是生物圈重要组分之一。由于全球气候变化世界多地冰川加速消融,暴露原本被冰盖覆盖的区域,这些区域被称为冰川消退区域(glacier retreat area)或冰川前部区域(glacier foreland)。自暴露开始消退区随即发生初生演替,随着演替进行,物质循环逐步建立,生物量和土壤C、N总量逐步增加。生态系统C、N输入最初以矿化外来物为主,逐渐转变为以生物固C、固N为主。演替早期生态系统的发育主要受土壤C、N含量的限制,而演替后期的限制性营养物转变为P。演替区域土壤逐渐发育并促进生态位的分化,细菌、真菌、古菌,病毒及其他微生物群落的生物量和多样性不断增加直至达到该地区可承受的极值。随着生存条件的改善,不同生态策略物种的更替导致每个演替阶段微生物群落结构的差异。整体上,伴随演替进行微生物群落丰度、结构和活性呈现梯度性变化。气候变化对冰川消退带生态演替结果产生多方面的影响,而这些影响结果又综合反馈气候变化,因此目前难以准确估计气候变化对消退带生态演替的净效应。综述了近年冰川消退带微生物群落演替方面相关的研究结果,同时分别对该区域物质循环的建立、微生物群落演替和气候变化造成的影响这三个方面进行详细描述,并指出当前研究的不足。     

Inverse relationship of epilithic algae and pelagic phosphorus in unproductive lakes: Roles of N2 fixers and light

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Bacterial Community Succession in Natural River Biofilm Assemblages

Temporal bacterial community changes in river biofilms were studied using 16S rRNA gene-based polymerase chain reaction–denaturing gradient gel electrophoresis (DGGE) followed by sequence analysis. Naturally occurring biofilms were sampled in 2001 during an undisturbed 7-month low-water period in the River Garonne (SW France). During the sampling period epilithic biomass exhibited a particular pattern: two 3-month periods of accumulation that resulted in two peaks in summer and fall, each at about 25 g ash-free dry mass per square meter. Bacterial community DGGE profiles differed between the summer and fall biomass peaks and shared only 30% common operational taxonomic units (OTUs), suggesting the influence of seasonal factors on these communities. During the second biomass accrual phase, bacterial richness and the appearance of new OTUs fitted a conceptual model of bacterial biofilm succession. During succession, five OTUs (corresponding to Dechloromonas sp., Nitrospira sp., and three different Spirosoma spp.) exhibited particular patterns and were present only during clearly defined successional stages, suggesting differences in life-history strategies for epilithic bacteria. Co-inertia analysis of DGGE banding patterns and physical–chemical data showed a significant relationship between community structure and environmental conditions suggesting that bacterial communities were mainly influenced by seasonal changes (temperature, light) and hydrodynamic stability. Within the periods of stability, analysis of environmental variables and community patterns showed the dominant influence of time and maturation on bacterial community structure. Thus, succession in these naturally occurring epilithic biofilm assemblages appears to occur through a combination of allogenic (seasonal) and autogenic changes.     

Secondary succession and effects of clear-logging on diversity in the subtropical forests on Okinawa Island,southern Japan

In order to clarify the recovery process of the subtropical forest on Okinawa Island, southern Japan, biomass accumulation and the successional trend of species diversity with time were investigated by comparing plots of old-growth and clear logged secondary forests. Self-thinning was an important factor in the development of young secondary forests, and the small variance in tree size within a stand was related to the stand not being fully stratified after clear-cutting. A large variance of size structure in old secondary and old-growth forests implies re-initiation of the understorey. Additionally, the trajectory of stand development indicated that the subtropical forest quickly recovered aboveground biomass, which reached its upper limit at about 50 years after disturbance. However, there was a large distinction in species diversity between the secondary forests and old-growth forests. The diversity of forest floor plants did not recover well after being clear-cut. This indicates that management of the subtropical forest should not only take timber-oriented tree species into account, but also the biodiversity in ground flora. The secondary forests were characterized by Castanopsis sieboldii and Schima wallichii, and the monopolization of C. sieboldii through secondary succession had a negative influence on species diversity. Distylium racemosum dominated at the late development stage and was considered a long-lived competing species that reduced the dominance of C. sieboldii and facilitated the co-occurrence of understorey species. Light-demanding pioneer tree species such as S. wallichii that regenerated after logging should be replaced by competitive effects of climax species, and thus relayed floristic change might increase species diversity along secondary succession.Nomenclature: Hatushima and Amano (1994).     

Seasonal succession of phytoplankton and its diversity in two highly eutrophic lakes with different conditions of stratification

Seasonal succession of phytoplankton biomass, its diversity and its photosynthetic activity in two highly eutrophic lakes have been compared. In order to test the intermediate disturbance hypothesis, the lakes have been chosen with almost the same level of trophy but different conditions of stratification, through two ice-free periods of open water with different weather conditions.High phytoplankton diversity throughout the period of investigation was characteristic for the shallower Lake Lobardzu. The number of species here was usually more than 30 and the Shannon diversity changed from 1.2 to 4.2. Owing to the frequent external disturbances, periods characterized by autogenic succession with establishing dominance and declining diversity alternated with periods of biomass reduction and rises of diversity and photosynthetic activity. In the warmer summer of 1983, with more intense warming of bottom layers and predominance of blue-greens, phytoplankton biomass was higher and diversity lower than in the cold summer of 1982.In stratified Lake Rudusku, phytoplankton diversity and number of species were usually much lower. During the long summer stratification up to three-four dominant species of blue-greens and dinoflagellates become established and competitive exclusion leading to low diversity advanced. Some changes in biomass and diversity, were caused by zooplankton activity.     

海南岛热带山地雨林天然次生林的功能群划分

热带林极高的物种丰富度使许多生态分析非常困难,把功能相似的物种划分为不同功能群,将为热带林的生态研究提供新的途径。以物种的7个功能特性因子(生长型、分布的海拔高度、分布的林型、木材密度、喜光性、演替地位和寿命)和9个林分结构因子(相对生物量、相对胸高断面积、相对树高、相对密度、相对频度、相对冠幅、相对更新数、相对死亡数和相对萌生数)为基础,应用数量化分析的方法,对海南岛典型的热带山地雨林天然次生林群落进行了功能群的划分。结果表明:(1)应用CCA分析林分结构因子时,可将山地雨林天然次生林的物种划分为6类功能群,它们的相对生物量、相对密度、相对频度、相对更新数、相对萌生数和相对死亡数等,随胸径和高度的增加而呈现有规律的变化;(2)应用CCA分析物种功能特性因子时,可将山地雨林天然次生林的物种划分为5类功能群,它们的功能特性都随演替过程而呈现有规律的变化;(3)在综合考虑两个不同角度CCA分析的基础上,最后将热带山地雨林天然次生林的物种共划分为11类功能群,它们能充分体现物种随胸径和高度结构的变化特点及其在演替过程中所处的阶段;(4)演替初期的灌木类功能群与各不同演替时期的乔木功能群共同分布于的中下层,但其大多处于死亡状态;(5)演替初期与演替中后期的乔木功能群则共同组成的主层林,但其死亡数量也较高。可见,海南岛热带山地雨林天然次生林目前正处不同功能群的激烈竞争阶段。