Tree diversity promotes predator but not omnivore ants in a subtropical Chinese forest

1. Epigeic ants are functionally important arthropods in tropical and subtropical forests, particularly by acting as predators. High predation pressure has been hypothesised to be a mechanism facilitating high diversity across trophic levels. 2. In this study, standardised pitfall traps were used in a highly diverse subtropical forest to test if and how ant species richness is related to tree species richness and a comprehensive set of other environmental variables such as successional age, soil properties or elevation. 3. A total of 13 441 ant individuals belonging to 3839 species occurrences and 71 species were collected, of which 26 species were exclusive predators and 45 species were omnivores. 4. Occurrence and species richness of total and omnivore ants were positively related to soil pH. Predator ant occurrence was unrelated to all environmental variables tested. 5. The species richness of predator ants increased with tree species richness but decreased with leaf functional diversity and shrub cover. Elevation negatively influenced only total ant species richness. 6. The evenness of predators increased with tree species richness, while the evenness of all ants decreased with shrub cover. Omnivore ant evenness decreased with tree evenness, but increased with successional age. 7. The results highlight the value of diverse forests in maintaining species richness and community evenness of a functionally important predator group. Moreover, the results stress the importance of analysing trophic groups separately when investigating biodiversity effects.     

Stand structure and woody species diversity in relation to stand stratification in a subtropical evergreen broadleaf forest, Okinawa Island

Stand structure and woody species diversity in a subtropical evergreen broadleaf forest grown in a silicate habitat, Okinawa Island, have been investigated on the basis of stand stratification. The forest stand consisted of four layers. The floristic composition of the top and the lower three layers was only slightly similar, although approximately one-third of the species were common to them. Mean tree weight decreased from the top toward the bottom layer whereas tree density increased from the top downward. This trend resembled the mean weight–density trajectory of self-thinning plant populations. The relationship between mean tree height and tree density for the upper two layers supported Yamakuras quasi –1/2 power law of tree height. The values of the Shannon–Wiener index, H, and the equitability index, J, tended to increase from the top layer downward except for the bottom layer. The values of H and J were, respectively, 4.83 bit and 0.82 for trees taller than 0.10 m. The lower layers contained many species of smaller height. High species diversity of the forest depended on small trees in the lower layers. Conservation of small trees in the lower layers, especially the bottom layer, is indispensable for sound maintenance of Okinawan evergreen broadleaf forests.     

冠层垂直高度对植物叶片功能性状的影响: 以鼎湖山南亚热带常绿阔叶林为例

植物功能性状能够反映植物对环境变化的响应, 从而影响生态系统功能。叶片是植物功能性状的重要研究对象, 但随着树冠垂直高度的增加, 植物叶片功能性状的变化规律仍不清楚。为了更好地解释植物对所处环境的响应机制, 本研究以鼎湖山南亚热带常绿阔叶林6个优势树种为研究对象, 对这些树种不同冠层高度的叶片比叶面积(SLA)、叶绿素相对含量(SPAD)、叶干物质含量(LDMC)、叶片碳含量(LCC)、叶片氮含量(LNC)、叶片磷含量(LPC)、叶片氮磷比(N : P)、等效水厚度(EWT)、氮平衡指数(NBI)、类黄酮含量(FLAV)共10个功能性状指标进行比较分析。结果表明: (1)随树冠垂直高度的增加, 叶干物质含量、类黄酮含量、等效水厚度和叶片磷含量呈增加趋势, 比叶面积和氮平衡指数呈减小趋势; 在冠层垂直方向上, 类黄酮含量与氮平衡指数、比叶面积与等效水厚度呈极显著负相关, 叶片氮含量与叶片磷含量呈极显著正相关。(2)不同树种叶片功能性状差异显著, 可能受植物自身特性和群落树种演替进程的影响。(3)叶片功能性状的种内差异沿垂直层次呈现不同的变化趋势。其中, 67%的树种的类黄酮含量随树冠垂直高度增加而显著增加, 67%的树种的氮平衡指数随树冠垂直高度增加而显著减小。南亚热带常绿阔叶林不同树种叶片功能性状随树冠垂直高度增加产生的变化, 反映了植物对光环境的适应以及对水资源和能量利用的不同策略。     

Tree diversity promotes predatory wasps and parasitoids but not pollinator bees in a subtropical experimental forest

From regional to global scales, anthropogenic environmental change is causing biodiversity loss and reducing ecosystem functionality. Previous studies have investigated the relationship between plant diversity and functional insect communities in temperate and also in tropical grasslands and forests. However, few studies have explored these dynamics in subtropical forests. Here, cavity-nesting Hymenoptera and associated parasitoids were collected across a controlled tree diversity experiment in subtropical China to test how predatory wasps, bees and parasitoids respond to tree species richness. Abundance and species richness of predatory wasps and parasitoids were positively correlated with tree species richness, while bee abundance and bee species richness were unrelated to tree species richness. Our results indicate that tree species richness increases the abundance and species richness of important communities such as predators and parasitoids. Moreover, the results highlight the importance of subtropical forests in maintaining abundance and species richness of key functional insect groups.     

鼎湖山南亚热带常绿阔叶林不同成熟度群落特征的比较

研究不同成熟度群落物种的共存状态,了解群落演替过程中结构的动态变化,对揭示森林不同发育时期的群落构建机制具有重要意义。本研究基于鼎湖山南亚热带常绿阔叶林20 ha固定监测样地(以下简称鼎湖山大样地)2010年植被调查的数据,根据径级与林龄将鼎湖山大样地划分为成熟林和过熟林两个斑块,对其群落物种组成、区系特征及径级结构进行了初步的比较。研究发现:(1)成熟林和过熟林种类组成的群落相似性系数高、物种多样性指数相近,以及锥栗(Castanopsis chinensis)重要值相似,但物种数、密度、总胸高断面积、物种重要值等有所不同;(2)热带区系成分在两个斑块均占优势,但以过熟林的比例较高;(3)两个斑块虽然所有活立木径级结构分布均呈倒"J"型,但在同径级个体群重要值比较中,成熟林中径级个体群重要值较高,而过熟林则是大径级个体群重要值较高。由此可见,鼎湖山大样地的成熟林与过熟林虽属于同一类群落的演替顶极阶段,但群落结构上却存在差异,推测在林窗作用下不同发育时期的群落存在不同的构建过程。     

Reprint of: Tree diversity promotes predatory wasps and parasitoids but not pollinator bees in a subtropical experimental forest

From regional to global scales, anthropogenic environmental change is causing biodiversity loss and reducing ecosystem functionality. Previous studies have investigated the relationship between plant diversity and functional insect communities in temperate and also in tropical grasslands and forests. However, few studies have explored these dynamics in subtropical forests. Here, cavity-nesting Hymenoptera and associated parasitoids were collected across a controlled tree diversity experiment in subtropical China to test how predatory wasps, bees and parasitoids respond to tree species richness. Abundance and species richness of predatory wasps and parasitoids were positively correlated with tree species richness, while bee abundance and bee species richness were unrelated to tree species richness. Our results indicate that tree species richness increases the abundance and species richness of important communities such as predators and parasitoids. Moreover, the results highlight the importance of subtropical forests in maintaining abundance and species richness of key functional insect groups.     

Density dependence is not very prevalent in a heterogeneous subtropical forest

There is a growing body of evidence demonstrating that tree survival is influenced by negative density‐dependence, but it is still controversial how the effect may vary with life‐stage, and to what extent it plays a role in regulating tree survival in heterogeneous subtropical forests. In this study, we investigated density‐dependent effects on tree survival of six tree species in a 5‐ha subtropical forest in eastern China. The roughly 45 000 individuals in the forest were fully censused in 2003 and 2008. For each of these species, we used an inhomogeneous pair‐correlation function to quantify the change in spatial distribution for different size classes, and a case‐control design to study seedling–adult associations in 2003. Autologistic regression was used to determine the influence of neighborhood factors on individual survival from 2003 to 2008. We found that seedlings of five species were repulsed by distance to nearest conspecific adults in terms of their survival, consistent with predictions of the Janzen–Connell mechanism. By contrast, only the least shade‐tolerant Schima superba had a negative relationship with individual survival and conspecific distance‐weighted basal area. This suggests that the Janzen–Connell effect is only prevalent at the early seedling stage or seed‐to‐seedling phase. The strength of clustering significantly declined at sapling–pole and pole–adult transitions for Sycopsis sinensis and at seedling–sapling transition for Cleyera pachyphylla. Correlations between individual survival and conspecific abundance for these species were consistent with trends in the strength of clustering. These results suggest that density dependence plays a limited role in individual survival and species spatial structure beyond the early seedling stage (i.e. after true leaves growing) in this forest. In addition, this study indicates that including individuals from early life‐stages and factoring out potential confounding factors such as habitat preference are important in studies that seek evidence for density dependence in forest trees.     

Stream salinization is associated with reduced taxonomic,but not functional diversity in a riparian plant community

Abstract Dryland salinity presents an overwhelming threat to terrestrial and aquatic habitats in Australia, and yet there remains very little empirical evidence of the impacts of secondary salinization on the biodiversity of riparian communities. Here we describe the response of a riparian plant community to stream and soil salinization, 25 years after the experimental clearing of a catchment in south‐western Australia. Riparian plant species diversity was inversely related to soil salinity, and plant species composition was significantly altered by increased soil salinity. Despite the evidence for an impact of salinization on the taxonomic diversity and composition of the riparian plant community, there was little evidence for any effect of salinization on functional group diversity, or on ecological functioning, as measured by the percentage of above‐ground plant cover.     

亚热带次生常绿阔叶林植物功能性状及多样性对人工抚育的响应

亚热带次生常绿阔叶林普遍存在生长滞缓、固碳功能和经济价值低下等问题,迫切需要通过人工抚育提高其生态系统功能。对植物功能性状及其多样性的深入研究,能够找寻植物共存与群落稳定发展的内在机制,为提升森林生态系统功能的实践提供基础。为了探究人工抚育对亚热带次生常绿阔叶林质量提升的影响,以杭州午潮山国家森林公园内的次生常绿阔叶林为研究对象,实施了不同强度的抚育措施,通过群落植被复查,分析群落植物功能性状以及功能多样性对不同强度人工抚育的响应。结果发现:抚育2年后,群落最大潜在树高与叶片氮、磷、钾含量显著上升,而叶片干物质含量和叶片叶绿素含量均有显著下降,比叶面积有下降趋势,两种强度的抚育之间差异不显著;叶片干物质含量、叶绿素含量与叶片氮含量间的相关性由不显著变为了显著负相关,而叶片干物质含量与叶片叶绿素含量,叶片磷、钾含量与叶片氮含量,叶片磷含量与叶片钾含量间的相关性由不显著变为显著正相关;在20%强度抚育下,功能丰富度指数均值从对照的0.52显著下降为0.16,功能均匀度指数与功能分离度指数的均值则分别上升到0.61和0.67,显著高于对照组的0.51和0.56;在10%强度抚育下,功能丰富度有所下降,功能均匀度与功能分离度均有所上升,但差异不显著。研究表明,人工抚育在一定程度上改变了群落水平的植物功能性状与功能多样性,提高了群落植物利用资源的能力和效率;而适度的抚育强度更能解放资源空间,对群落产生的影响更加明显。该结果为亚热带次生常绿阔叶林的经营与管理提供依据与参考。     

Ecological stoichiometry of the epiphyte community in a subtropical forest canopy

Epiphytes in tree canopies make a considerable contribution to the species diversity, aboveground biomass, and nutrient pools in forest ecosystems. However, the nutrient status of epiphytes and their possible adaptations to nutrient deficiencies in the forest canopy remain unclear. Therefore, we analyzed the stoichiometry of five macroelements (C, N, P, K, and Ca) in four taxonomic groups (lichens, bryophytes, ferns, and spermatophytes) to investigate this issue in a subtropical montane moist evergreen broad‐leaved forest in Southwest China. We found that the interspecific variations in element concentrations and mass ratios were generally greater than the intraspecific variations. And there were significant stoichiometric differences among functional groups. Allometric relationships between N and P across the epiphyte community indicated that P might be in greater demand than N with an increase in nutrients. Although canopy nutrients were deficient, most epiphytes could still maintain high N and P concentrations and low N:P ratios. Moreover, ferns and spermatophytes allocated more limited nutrients to leaves than to stems and roots. To alleviate frequent drought stress in the forest canopy, vascular epiphytes maintained several times higher K concentrations in their leaves than in the tissues of lichens and bryophytes. Our results suggest that epiphytes may have evolved specific nutrient characteristics and adaptations, so that they can distribute in heterogeneous canopy habitats and maintain the stability of nutrient metabolism.     

Host plant genus-level diversity is the best predictor of ectomycorrhizal fungal diversity in a Chinese subtropical forest

Microbial diversity is generally far higher than plant diversity, but the relationship between microbial diversity and plant diversity remains enigmatic. To shed light on this problem, we examined the diversity of a key guild of root-associated microbes, that is, ectomycorrhizal (EM) fungi along a plant diversity gradient in a Chinese subtropical forest. The results indicated that EM fungal diversity was positively correlated with host plant diversity. Furthermore, this relationship was best predicted by host genus-level diversity, rather than species-level diversity or family-level diversity. The generality of this finding was extended beyond our study system through the analyses of 100 additional studies of EM fungal communities from tropical and temperate forests. Here as well, EM fungal lineage composition was significantly affected by EM plant diversity levels, and some EM fungal lineages were co-associated with some host plant genera. These results suggest a general diversity maintenance mechanism for host-specific microbes based on higher order host plant phylogenetic diversity.     

Understorey versus canopy: patterns of vertical stratification and diversity among Lepidoptera in a Bornean rain forest

We studied the vertical distribution of Lepidoptera from a canopy walkway within a dipterocarp rain forest at Kinabalu Park (Borneo) using three different methods: (1) Bait traps to survey fruit-feeding nymphalid butterflies, (2) standardized counts for predominantly flower-visiting butterflies and their potential predators, aerial-hawking birds, and (3) attraction by blacklight for hawk- and tiger moths. There was a distinct decrease in the abundance of fruit-feeding nymphalids towards the canopy, probably due to a reduced and less predictable availability of rotting fruits in higher strata. These constraints might also be responsible for a higher abundance variation in the canopy, and a significant shift in size from larger species in the understorey to smaller ones in the canopy. Changes of microclimate and the conspicuous increase of insectivorous aerial-hawking birds from ground to canopy layer may be responsible for the prominent change in species composition of fruit-feeding nymphalids between 20 and 30 m. Nectar-feeding Lepidoptera showed a reversed abundance pattern. One main factor contributing to the much higher abundance of flower-visiting butterflies and moth taxa in the canopy, such as Sphingidae and some Arctiinae, might be the increase of nectar resources available in upper vegetation layers. A distinctly higher diversity in hawkmoths was also found in the canopy. A higher abundance of insectivorous aerial-hawking birds in the canopy might contribute to the shift in body design of fruit-feeding nymphalids from more slender bodies at lower vegetation layers to stouter ones (i.e. species which are stronger on the wing) in the canopy. Larval resources could play an additional role in specialisation on but a small part of the vertical gradient. This may explain stratification pattern of the nymphalid subfamilies Morphinae and Satyrinae. Monocotyledoneous larval food plants of both taxa, whose flight activity is largely restricted to the understorey, occur mostly in lower vegetation layers. Our observations on a wide taxonomic and ecological range of butterflies and moths indicate that tropical forest canopies hold a distinct and unique Lepidoptera fauna, whose species richness and abundance patterns differ from lower strata. However, the notion of tropical forest canopies as peaks of terrestrial diversity does not hold uniformly for all taxa or guilds.     

EcM fungal community structure, but not diversity, altered in a Pb-contaminated shooting range in a boreal coniferous forest site in Southern Finland

Boreal forests contain diverse fungal communities that form essential ectomycorrhizal symbioses with trees. To determine the effects of lead (Pb) contamination on ectomycorrhizal fungal communities associated with the dominant pine (Pinus sylvestris L.), we surveyed sporocarps for 3 years, analyzed morphotyped ectomycorrhizal root tips by direct sequencing, and 454-sequenced fungal communities that grew into in-growth bags during a 2-year incubation at a shooting range where sectors vary in the Pb load. We recorded a total of 32 ectomycorrhizal fungi that formed conspicuous sporocarps, 27 ectomycorrhizal fungal phylotypes from 294 root tips, and 116 ectomycorrhizal fungal operation taxonomic unit (OTUs) from a total of 8194 internal transcribed spacer-2 454 sequences. Our ordination analyses by nonparametric multidimensional scaling (NMS) indicated that the Pb enrichment induced a shift in the ectomycorrhizal community composition. This was visible as indicative trends in the sporocarp and root tip data sets, but was explicitly clear in the communities observed in the in-growth bags. The compositional shift in the ectomycorrhizal community was mainly attributable to an increase in the frequencies of OTUs assigned to genus Thelephora and to a decrease in the OTUs assigned to Pseudotomentella, Suillus, and Tylospora in Pb-contaminated areas when compared with the control. While the compositional shifts are clear, their functional consequences for the dominant trees or soil ecosystem function remain undetermined.     

Genetic diversity affects productivity in early but not late stages of stand development

Recent declines in the genetic diversity of populations have stimulated research on the importance of genetic diversity for the functioning of natural communities. Current studies on this topic are based on the exploration of a limited number of clones and do not allow distinctions to be made between the effects of genetic identity and genetic diversity per se and to evaluate the effects of genetic diversity in genetically diverse communities. Also, most information comes from short-term studies, which are insufficient for evaluating the long-term effects relevant in relatively undisturbed communities of perennial species.We explored the importance of clone diversity vs. clone identity for stand productivity and the changes of the pattern over time. We used 18 clones of a perennial grass, Festuca rubra, to establish a set of communities composed of 1, 6 or 18 clones in two environments and studied the effects of genetic diversity on stand productivity over 3 years.Genetic diversity had a significant effect on stand productivity in the 1st year but not in the 2nd or 3rd year. In most cases, the observed yield was not significantly different from the total expected yield. The biomass of the mixtures never outperformed the biomass of the most productive clone, suggesting that clone identity is an important determinant of total biomass.The results indicate that the effects of genetic diversity on stand productivity may be transient and suggest that the conclusions of short-term studies on diversity effects should be evaluated carefully. They also suggest that individual clones are not complementary and that the properties of the stands are mainly additive results of the properties of the constituent clones.     

Spatial structure and diversity of woody plants and ectomycorrhizal fungus sporocarps in a natural subtropical forest

Spatial patterns of ectomycorrhizal fungi, ectomycorrhizal plants, and non-ectomycorrhizal plants were investigated in a natural subtropical forest using second-order analysis. The results of spatial pattern analysis showed that the degree of clumping of woody plants and ectomycorrhizal sporocarps were correlated. There was a significantly positive correlation of relative aggregation indices between ectomycorrhizal fungi and both non-ectomycorrhizal trees and ectomycorrhizal saplings. Correlations between percentage of ectomycorrhizal trees and sporocarp occurrence of ectomycorrhizal fungi and between diversities of woody plants and ectomycorrhizal fungi were distance-dependent or scale-related. A significantly high percentage of ectomycorrhizal trees was found only at relatively short distance from ectomycorrhizal fungal sporocarps, and significantly positive correlation of the diversity between woody plants and ectomycorrhizal fungi was found only at relative long distance, which implied that ectomycorrhizal sporocarps prefer ectomycorrhizal-tree-dominant micro-sites at near distances and at relatively large scales, diverse ectomycorrhizal sporocarps could be found in woodlands with high diversity of woody plants. Important factors affecting the spatial distribution, occurrence, and diversity of ectomycorrhizal fungi include spatial pattern of ectomycorrhizal plants and non-ectomycorrhizal plants, percentage of ectomycorrhizal plants, and plant diversity in a natural forest.     

Physical,but not chemical,antiherbivore defense expression is related to the clustered spatial distribution of tropical trees in an Amazonian forest

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Considerable MHC diversity suggests that the functional extinction of baiji is not related to population genetic collapse

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