Species mixture effects on flammability across plant phylogeny: the importance of litter particle size and the special role for non‐Pinus Pinaceae

Fire affects and is affected by plants. Vegetation varies in flammability, that is, its general ability to burn, at different levels of ecological organization. To scale from individual plant traits to community flammability states, understanding trait effects on species flammability variation and their interaction is important. Plant traits are the cumulative result of evolution and they show, to differing extents, phylogenetic conservatism. We asked whether phylogenetic distance between species predicts species mixture effects on litterbed flammability. We conducted controlled laboratory burns for 34 phylogenetically wide‐ranging species and 34 random two‐species mixtures from them. Generally, phylogenetic distance did not predict species mixture effects on flammability. Across the plant phylogeny, most species were flammable except those in the non‐Pinus Pinaceae, which shed small needles producing dense, poorly ventilated litterbeds above the packing threshold and therefore nonflammable. Consistently, either positive or negative dominance effects on flammability of certain flammable or those non‐flammable species were found in mixtures involving the non‐Pinus Pinaceae. We demonstrate litter particle size is key to explaining species nonadditivity in fuelbed flammability. The potential of certain species to influence fire disproportionately to their abundance might increase the positive feedback effects of plant flammability on community flammability state if flammable species are favored by fire.     

The biogeography and filtering of woody plant functional diversity in North and South America

Aim In recent years evidence has accumulated that plant species are differentially sorted from regional assemblages into local assemblages along local‐scale environmental gradients on the basis of their function and abiotic filtering. The favourability hypothesis in biogeography proposes that in climatically difficult regions abiotic filtering should produce a regional assemblage that is less functionally diverse than that expected given the species richness and the global pool of traits. Thus it seems likely that differential filtering of plant traits along local‐scale gradients may scale up to explain the distribution, diversity and filtering of plant traits in regional‐scale assemblages across continents. The present work aims to address this prediction. Location North and South America. Methods We combine a dataset comprising over 5.5 million georeferenced plant occurrence records with several large plant functional trait databases in order to: (1) quantify how several critical traits associated with plant performance and ecology vary across environmental gradients; and (2) provide the first test of whether the woody plants found within 1° and 5° map grid cells are more or less functionally diverse than expected, given their species richness, across broad gradients. Results The results show that, for many of the traits studied, the overall distribution of functional traits in tropical regions often exceeds the expectations of random sampling given the species richness. Conversely, temperate regions often had narrower functional trait distributions than their smaller species pools would suggest. Main conclusion The results show that the overall distribution of function does increase towards the equator, but the functional diversity within regional‐scale tropical assemblages is higher than that expected given their species richness. These results are consistent with the hypothesis that abiotic filtering constrains the overall distribution of function in temperate assemblages, but tropical assemblages are not as tightly constrained.     

Temporal changes in the dominance of tree functional traits,but no changes in species diversity during woody plant encroachment in a Brazilian savanna

Questions : Woody encroachment in savannas has been associated with changing taxonomic composition and ecosystem function. Interestingly, there is little understanding of how encroachment impacts plant functional diversity and how those changes relate to plant demography, a crucial mediator between taxonomic composition and ecosystem function. Location : Southeastern Brazil. Methods: Using a landscape scale fire suppression experiment in a diverse Brazilian savanna, we quantify how change in species composition over seven years impacted vegetative and reproductive tree functional diversity as determined by new recruits, dead and surviving trees. Results: Over seven years, tree above-ground biomass increased by 15%, while total species richness did not change. Despite minor changes, species composition remained overall similar (82%), with few species contributing significantly to plot dissimilarity over time. There were small changes in vegetative traits, where the community-weighted mean increased in maximum tree height (↑ 2.1%) and specific leaf area (↑ 5.3%), and decreased in wood density (↓ 1.3%) and bark thickness (↓ 9.4%). Changes in reproductive traits were larger than in vegetative traits, with an increase in the prevalence of monoecy (↑ 32.6%), dioecy (↑ 44.2%), large seeds (↑ 20.3%), animal-mediated seed dispersal (↑ 4.9%) and pollination by very small insects (↑ 45.5%), and a decrease in the prevalence of hermaphroditism (↓ 9%), small seeds (6.8%) and pollination by small insects (12.5%). The overall decrease in bark thickness and increase in monoecy and dioecy were mainly driven by characters of the new recruits, while the overall increase in specific leaf area (SLA) and decrease in small seeds appeared largely determined by the loss of trees possessing those traits. Conclusions: Encroachment leads to changes that are likely increasing ecosystem vulnerability to fire and drought. Further, the compositional changes observed appear to drive marked change in reproductive traits, indicating increasing dependence on animals for dispersal and reproduction. Understanding post-hoc encroachment impacts in an era of widespread pervasive encroachment is fundamental to reconciling ecosystem functions such as nutrient cycling and pollination services as there is a loss of species with open ecosystem life-history strategies. Among savannas, there remains an urgent need to understand relationships between woody cover and ecosystem function to determine thresholds in woody cover promoting resilient savanna ecosystems.     

The resprouting response of co‐occurring temperate woody plant and grass species to elevated [CO2]: An insight into woody plant encroachment of grasslands

The phenomenon of woody plant thickening in grasslands has been observed globally and is likely to have widespread ecological consequences. It has been proposed that woody plant thickening is driven in part by rising atmospheric [CO₂] enhancing the resprouting ability of woody plants relative to grasses so they respond more strongly after disturbances such as herbivory and fire. The aim of this study was to examine the CO₂ effect on the resprouting ability of 16 co‐occurring temperate woody plant and grass species (eight species from each growth form). Plants were grown in a controlled glasshouse experiment under ambient (400 ppm) and elevated [CO₂] (600 ppm) for 14 weeks after which their resprouting ability was measured. Root non‐structural carbohydrate (NSC_mass) and nitrogen (N_mass) storage was used as proxies to measure the resprouting ability of woody plants while for the grasses it was measured directly. We found that both the woody plants (22% on average; P = 0.003) and grasses (20% on average; P = 0.003) produced more biomass under elevated [CO₂]. Despite the woody plants not allocating additional carbon to belowground storage under elevated [CO₂], they had significantly greater root NSC_mass (23% on average; P = 0.007) due to increased root biomass production (8% on average; P = 0.007). In contrast, root N_mass of the woody plants did not differ between CO₂ treatments (P = 0.373). Surprisingly, the resprouting ability of the grasses did not significantly differ between the CO₂ treatments (P = 0.067). These results provide evidence that the differing resprouting response of woody plants and grasses under elevated [CO₂] may be contributing to woody plant encroachment of grasslands worldwide.     

植物功能性状对土壤保持的影响研究述评

植被对土壤保持具有重要的影响,但是从植物功能性状的角度总结评述植被对土壤保持影响的研究并不多见。总结评述了植物地上功能性状、地下功能性状对土壤保持功能的影响以及植物地上、地下功能性状的关系,认为:(1)植被地上部分功能性状对土壤保持的作用主要体现在对溅蚀、面蚀的影响及间接改变土壤理化性质等方面,其功能性状指标主要包括叶面积、叶长、叶宽、枝数、植被高度等;(2)植被地下部分功能性状对土壤保持的作用主要体现在固持土壤、提高土壤抗剪切强度、提高土壤抗侵蚀能力、增强土壤渗透性,植物根系固持土壤与根系抗拉能力密切相关,植物根系土壤的物理和水文性质,与细根比例、根长密度、根表面积等性状密切相关;(3)可以通过植物地上部分功能性状间接反映地下部分功能性状,但是现有研究多为定性认识;(4)在植物功能性状对土壤保持的研究中亟待加强植被地上地下功能性状的长期定位监测,深化植被功能性状尤其是根系特征与土壤保持的作用机理,加强植被地上部分、地下部分功能性状的定量表达,建立植被功能性状与土壤保持功能的定量关系,实现植被功能性状与土壤保持功能特征的动态链接。     

Ecology and biogeography in 3D: The case of the Australian Proteaceae

The key biophysical pressures shaping the ecology and evolution of species can be broadly aggregated into three dimensions: environmental conditions, disturbance regimes and biotic interactions. The relative importance of each dimension varies over time and space, and in most cases multiple dimensions need to be addressed to adequately understand the habitat and functional traits of species at broad spatial and phylogenetic scales. However, it is currently common to consider only one or two selective pressures even when studying large clades. We illustrate the importance of the all‐inclusive multidimensional approach with reference to the large and iconic plant family, Proteaceae: we review life‐history traits related to these three dimensions for the 46 genera occurring in Australia and show that this family can be considered the product of a long history of harsh environments, recurrent fires and strong faunal interactions. Because most Proteaceae species occur in fire‐prone ecosystems and possess fire‐adaptive traits that are both ancient and essential for their survival, disturbance by fire is likely to explain much of this family's ecology, evolution and distribution. Approaches that only examine prevailing environmental variables may fail to identify the mechanisms that drive a taxon's biogeography; they need to consider the likely mechanisms of adaptation and accept or reject plausible alternative hypotheses as the evidence allows. As multidisciplinary teams that consider all aspects of a taxon's ecology are assembled, and databases and numerical tools become increasingly available, studies on the ecology, biogeography and diversity of organisms at broader spatial and phylogenetic scales will arrive at more realistic conclusions.     

践踏干扰对碧塔海高寒草甸植被茎叶性状、物种多样性和功能多样性的影响

高寒草甸具有重要的生态服务功能,然而固有脆弱性使其极易遭受气候变化和人为干扰等多重因素的影响。作为滇西北旅游资源中重要的组分之一,高寒草甸吸引了大批游客前往开展徒步旅行活动,但伴随着的践踏干扰作用会不可避免地对高寒草甸生态系统带来负面影响。然而,目前关注践踏干扰对滇西北高寒草甸植被的影响,特别是植被功能性状和功能多样性如何发生变化方面的研究还十分欠缺。以云南省香格里拉市碧塔海自然保护区内的典型高寒草甸生态系统为研究对象,采用实验践踏的方式(一共5种不同强度的践踏处理)来模拟旅游活动对草甸植被的干扰作用,并以草甸植被的茎叶性状特征为切入点,重点探讨践踏干扰对茎叶性状的平均大小和变异程度的影响,以及物种丰富度(以物种形态分类为基础)和功能丰富度(以功能性状为基础)之间的关系。研究结果显示,随着践踏强度的增加,植株高度和叶片大小的平均值,而不是茎叶性状的变异程度,出现明显下降趋势。此外,物种丰富度和功能丰富度均随践踏强度的增强而减小,且两者之间呈现显著正相关关系。然而,较之轻度践踏实验组,重度践踏实验组中的功能均匀度和功能分离度水平均有所增加,表明践踏干扰可能会在短期内打破优势种对资源的绝对占有格局和减少物种间的生态位重叠程度。尽管高寒草甸对人类践踏活动有一定的承受能力,但气候变化和人为干扰等多重因素势必会改变和影响高寒草甸群落的结构和功能可持续性,这也对高寒草甸的保护与管理工作提出了更加紧迫的要求。     

The impact of herbivore exclusion on forb diversity: Comparing species and functional responses during a drought

Rainfall and herbivory shape savannah herbaceous communities, but these disturbances are being altered globally. To assess potential consequences of such alterations, we evaluated herbivore effects on species and functional diversity during an episodic drought in a sodic savannah using data collected from long-term herbivore exclosures in the Kruger National Park, South Africa. Herbaceous life forms are rarely acknowledged as distinct functional entities. Moreover, the functional ecology of forbs remains elusive. Here, we present disturbances–responses by forbs separately from grasses. We hypothesised that combinations of intense utilisation and drought would be associated with low diversity and high dominance at species and functional levels for both life forms. Contrary to our hypothesis, low forb and grass diversity was associated with long-term herbivore exclusion, which exceeded expected undesirable effects of intense utilisation and drought. Grasses responded less sensitively, suggesting that forbs respond dynamically to changes in herbivore assemblage when these alterations are combined with drought. Consistent with patterns in savannah systems, forbs contributed significantly to species and functional trait diversity. High forb diversity is suggested to enhance resilience of this nutrient-rich ecosystem against declines in its functioning when subjected to drought and alterations in herbivory.     

Biological significance of dead biomass retention trait in Mediterranean Basin species: an analysis between different successional niches and regeneration strategies as functional groups

Standing dead biomass retention is considered one of the most relevant fuel structural traits to affect plant flammability. However, very little is known about the biological significance of this trait and its distribution between different functional groups. Our aim was to analyse how the proportion of dead biomass produced in Mediterranean species is related to the successional niche of species (early‐, mid‐ and late‐successional stages) and the regeneration strategy of species (seeders and resprouters). We evaluated biomass distribution by size classes and standing dead biomass retention in nine dominant species from the Mediterranean Basin in different development stages (5, 9, 14 and 26 years since the last fire). The results revealed significant differences in the standing dead biomass retention of species that presented a distinct successional niche or regeneration strategy. These differences were restricted to the oldest ages studied (>9 years). Tree and small tree resprouters, typical in late‐successional stages, presented slight variations with age and a less marked trend to retain dead biomass, while seeder shrubs and dwarf shrubs, characteristic of early‐successional stages, showed high dead biomass loads. Our results suggest that the species that tend to retain more dead branches are colonising species that may promote fire in early‐successional stages.     

漓江河岸带木本植物功能性状跨尺度变异研究

研究植物功能性状变异,有助于揭示植物对环境的适应策略以及群落构建机制。以漓江河岸带木本植物为研究对象,利用广义线性模型和"varcomp"函数对漓江河岸带主要木本植物的3个叶功能性状（比叶面积SLA;叶面积LA;叶厚度LTH）跨不同尺度（样方、物种、个体、叶）变异进行方差分解,分析各尺度的相对贡献及变异的主要来源。结果表明：（1）植物功能性状在4个巢式尺度的解释方差规律较一致,均表现为物种（0.51-0.70） > 个体（0.21-0.22） > 叶和残差（0.09-0.20） > 样方（0-0.07）;（2）4个功能性状种间尺度的解释方差均最大（SLA、LA和LTH分别为64%、70%和51%）;其次种内尺度的解释方差（SLA、LA和LTH分别为34%、30%和42%）仅次于种间。（3）样方尺度的解释方差很小甚至为零。发现：（1）植物功能性状变异主要由种间变异驱动;（2）种内性状变异是功能性状变异的重要且不容忽视的来源;（3）生境过滤并非直接作用于群落间物种组成,而可能是通过直接作用于植物功能性状的总体分布来影响植物群落构建过程,基于功能性状的生境过滤可能在漓江河岸带植物群落构建中起重要作用。     

高寒草甸15种植物种子发芽的比较研究

对高寒草甸 15种植物种子的发芽进行了比较实验研究。结果显示 ,冷湿层化、温度变幅及光照条件能够提高或者降低多数高寒草甸植物种子发芽率。其中 ,13种植物对层化、11种对光照条件、14种对温度变幅处理有显著性响应。 15种植物中 ,有14种对单一因子或因子组合有反应 ,仅藏嵩草种子发芽对设定的因子或因子组合没有响应。根据不同植物种子对不同处理及其组合的发芽反应可将植物种子划分为不同的反应类型 ,通过对种子进行冷湿层化处理 ,可以部分或者全部地替代某些植物种子发芽对光、温需求。探明植物种子在特定环境因子组合条件下的发芽表现 ,对通过种子恢复退化草甸是至关重要的。     

根系功能性状对干瘠立地适应的种间差异——以北京石质山地主要观赏树种为例

分析干瘠立地中不同树种根功能性状的变化及差异,研究不同树种对干瘠环境的适应对策。试验材料为林龄、林分密度相对一致的石质山区主要观赏树种（黄栌、山桃、栓皮栎）,采用挖掘法获取树木细根,测定细根形态参数。根据田间持水量和石砾含量分别设置4个干瘠梯度（轻度、中度、重度、极度）,对比分析北京干瘠地3种树种根功能性状,探讨干瘠立地中不同树种根系形态的差异及其对特殊生境的适应。结果表明：干旱、瘠薄均对树木的根功能性状产生影响,但对不同性状以及不同树种的影响程度具有差异,其中对山桃根系的影响最大;3种树种根功能性状在不同干瘠等级下存在显著变化,同一性状的变化因树种不同而异。从轻度到极度干旱,3种树种的RL均逐渐减小,但是黄栌变化不显著;黄栌、栓皮栎RTD随干瘠的加剧而减小,而山桃随干瘠的加剧而增加;黄栌和山桃SRL随干旱、瘠薄的加剧,表现出一致的规律,即从轻度到极度,其SRL随即减小,而栓皮栎SRL随干旱、瘠薄的加剧而增大;随干瘠的加剧,黄栌SRA显著减小,山桃、栓皮栎SRA随干瘠的加剧而增大。树木对干瘠环境的变化都有一定的响应能力及适应对策,能够通过自身的形态变化来适应不断变化的环境,但不同物种对干瘠环境变化的适应途径不同,通过权衡碳分配、碳消耗采取了相应的生态适应策略,不同树种间存在明显差异。本研究3种树种中,黄栌在干瘠环境下根系性状变化不显著,耐干瘠能力较强;干瘠胁迫对山桃根系性状影响显著,山桃通过主动调节根系形态变化来适应不断变化的环境,具有较强的抗干瘠能力;栓皮栎的细根数量多,吸收土壤水分、养分能力较强,能够在干瘠环境中快速生长。     

New insights into leaf and fine‐root trait relationships: implications of resource acquisition among 23 xerophytic woody species

Functional traits of leaves and fine root vary broadly among different species, but little is known about how these interspecific variations are coordinated between the two organs. This study aims to determine the interspecific relationships between corresponding leaf and fine‐root traits to better understand plant strategies of resource acquisition. SLA (Specific leaf area), SRL (specific root length), mass‐based N (nitrogen) and P (phosphorus) concentrations of leaves and fine roots, root system, and plant sizes were measured in 23 woody species grown together in a common garden setting. SLA and SRL exhibited a strong negative relationship. There were no significant relationships between corresponding leaf and fine‐root nutrient concentrations. The interspecific variations in plant height and biomass were tightly correlated with root system size characteristics, including root depth and total root length. These results demonstrate a coordinated plant size‐dependent variation between shoots and roots, but for efficiency, plant resource acquisition appears to be uncoupled between the leaves and fine roots. The different patterns of leaf and fine‐root traits suggest different strategies for resource acquisition between the two organs. This provides insights into the linkage between above‐ and belowground subsystems in carbon and nutrient economy.