常绿阔叶林外生和丛枝菌根树种细根形态和构型性状对氮添加的可塑性响应

以中亚热带常绿阔叶林外生菌根树种罗浮栲和丛枝菌根树种木荷为研究对象,采用根袋法进行野外原位氮添加试验,研究了细根形态性状(比根长、比表面积、组织密度、平均根直径)和构型性状(分枝数、分枝比、根长增长速率、根尖密度、分枝密度),分析不同菌根树种细根形态和构型性状对氮沉降的响应。结果表明: 随序级增加,外生和丛枝菌根树种细根比根长、比表面积和分枝数对氮添加的塑性响应逐渐降低,组织密度则相反;这反映了不同分枝等级细根的养分获取与资源维持在序级间存在权衡关系。不同菌根树种对土壤氮有效性的变化采取不同的适应对策: 氮添加后,罗浮栲细根采取机会主义策略,依靠细根本身来提高养分吸收效率、增强空间扩展和就地养分吸收能力,以快速的养分吸收策略为主;而木荷通过养分吸收效率和根系构建成本之间的权衡,并未改变细根形态性状,更多地依赖于菌根菌和细根构型之间的互补性进行养分获取。外生和丛枝菌根树种维持和构建细根碳(C)成本的差异,导致细根采取最适合自身的养分捕获方式,以达到生存的最优策略。     

亚热带典型树种根毛特征及其与共生真菌的关系

根毛和共生真菌增加了吸收面积,提高了植物获取磷等土壤资源的能力。由于野外原位观测根表微观结构较为困难,吸收细根、根毛、共生真菌如何相互作用并适应土壤资源供应,缺乏相应的数据和理论。该研究以受磷限制的亚热带森林为对象,选取了21种典型树种,定量了根毛存在情况、属性变异,分析了根毛形态特征与共生真菌侵染率、吸收细根功能属性之间的关系,探讨了根表结构对低磷土壤的响应和适应格局。结果表明:1)在亚热带森林根毛不是普遍存在的, 21个树种中仅发现7个树种存有根毛, 4个为丛枝菌根(AM)树种, 3个为外生菌根(ECM)树种。其中,马尾松(Pinus massoniana)根毛出现率最高,为86%;2)菌根类型是理解根-根毛-共生真菌关系的关键,AM树种根毛密度与共生真菌侵染率正相关,但ECM树种根毛直径与共生真菌侵染率负相关; 3) AM树种根毛长度和根毛直径、ECM树种根毛出现率与土壤有效磷含量呈负相关关系。该研究揭示了不同菌根类型树种根毛-共生真菌-根属性的格局及相互作用,为精细理解养分获取策略奠定了基础。     

中国亚热带不同菌根树种的根叶形态学性状特征与生长差异: 以江西新岗山为例

探究植物功能性状的种内和种间变异不仅有助于揭示植物对环境的适应, 也能够反映植物的生态策略, 但不同菌根类型树木生长过程中根叶形态学功能性状的适应策略仍有待探究。本研究依托中国亚热带森林生物多样性与生态系统功能实验研究平台(BEF-China)选取7种丛枝菌根(AM)树木和7种外生菌根(EM)树木的纯林, 测定各个树种的比叶面积、叶干物质含量、比根长、根系直径、树高生长速率、地径生长速率及细根生物量等根叶形态学功能性状和生长指标, 探讨了两种菌根类型树种间的根叶形态学特征的差异。结果表明: 与AM树种相比, EM树种具有较小的比叶面积、吸收根平均直径和生长速率, 但具有更大的叶干物质含量; 两种菌根树种之间的比根长和细根生物量无显著差异。比叶面积、叶干物质含量、树高生长速率、地径生长速率和细根生物量等功能性状及生长指标在不同菌根类型、树种及二者的交互作用中均存在显著差异; 且树种、根功能型、菌根类型及三者之间的交互作用均对根功能性状有显著影响。EM树种地上指标的种内变异均大于种间变异, 而AM树种地上指标的种内和种间变异程度类似; 但两种菌根树种细根生物量的种间变异均大于种内变异。尽管两种菌根树种地上部分生长速率较快通常表现为较低的叶干物质含量, 但AM树种通常拥有较高的吸收根比根长, 而EM树种拥有较粗的运输根平均直径。吸收根比根长越低, 两类菌根树种的细根生物量就越多。由此可见, 根叶功能性状对植物地上部分的生长具有一定的协同效应, 其中运输根主要在EM树种地上生长过程中发挥重要作用, 吸收根主要与AM树种的地上部分生长有关; 但两类菌根树种的地下细根生物量均与吸收根有关。     

滨海沙地主要造林树种叶片功能性状及养分重吸收特征

为探讨不同树种对滨海沙地干旱贫瘠环境的适应策略,以滨海沙地主要造林树种木麻黄、湿地松、厚荚相思和尾巨桉为对象,研究了不同树种叶片功能性状及养分重吸收特征.结果表明:阔叶树种(厚荚相思和尾巨桉)的叶面积、比叶面积显著高于针叶树种(木麻黄和湿地松),而针叶树叶干物质含量、叶厚度最高.成熟叶和凋落叶的N、P含量表现为阔叶树高于针叶树,成熟叶高于凋落叶,但凋落叶N∶P较高.针叶树种的N、P养分重吸收效率大于阔叶树种,P重吸收效率明显高于N,木麻黄、湿地松、厚荚相思和尾巨桉的N、P吸收效率分别为64.2%、63.1%、47.0%、16.8%和92.5%、81.6%、80.3%、18.0%.比叶面积与叶片N、P含量呈显著正相关,与叶干物质含量,叶厚度以及N、P养分重吸收效率呈显著负相关;叶干物质含量与叶厚度及N、P养分重吸收效率呈显著正相关.就叶片功能的权衡关系而言,木麻黄和湿地松属于缓慢投资-收益型物种,具有较高的养分重吸收效率,而厚荚相思和尾巨桉属于快速投资-收益型物种,养分的重吸收效率较低.不同滨海沙地造林树种通过叶片功能性状及养分重吸收之间的相互协调实现对滨海沙地特殊生境的适应性.     

菌根类型对亚热带森林主要树种根叶功能性状关联的影响

叶片和吸收根(即1级根)是植物地上和地下资源获取器官，研究二者功能性状的关系及其影响因素，有助于深入理解植物生长过程中对资源的利用和分配。本研究在江西省九连山国家自然保护区选择了10种丛枝菌根(AM)树种和8种外生菌根(EM)树种，分析了叶片和吸收根的形态性状(叶厚度、比叶面积、叶组织密度、根直径、比根长、根组织密度)和化学性状(叶与根的碳、氮浓度及碳氮比)在不同菌根类型间的变异及关联。结果表明：与AM树种相比，EM树种的叶片较厚、比叶面积较小，且根直径较细、根组织密度和根碳浓度及碳氮比较低。AM和EM树种的叶片与吸收根的形态性状均不相关，而化学性状(氮浓度和碳氮比)在AM树种中呈正相关，但在EM树种中则不相关。主成分分析结果表明，不管是AM还是EM树种，比叶面积与比根长均呈正交分布，可见叶片与吸收根获取资源具有独立性。此外，根直径与根氮浓度在所有树种和AM树种中均表现出显著正相关，而在EM树种则呈临界负相关。根直径与比根长则是在所有树种和不同菌根类型树种中均呈正相关。总之，本研究的结果强调了菌根类型在调节叶片与吸收根功能性状之间的关系上具有重要性，并建议今后在评估木本植物根系的不同功...     

不同林龄刺槐叶片养分重吸收特征及其对土壤养分有效性的响应

为揭示丘陵沟壑区刺槐的养分重吸收特征及其驱动因素,研究该区不同林龄刺槐叶片全氮和全磷的浓度,以及土壤有机碳、全氮、全磷、铵态氮、硝态氮和速效磷浓度及其化学计量,分析了叶片氮磷重吸收效率与土壤养分特性之间的关系。结果表明: 植物和土壤的养分随林龄增长发生显著变化,而土壤总磷和速效磷浓度较低。氮重吸收效率随林分生长先增加后降低,范围为48.2%～54.0%,平均为48.5%;磷重吸收效率则显著增加,范围为45.2%～49.4%,平均为46.9%。氮重吸收效率与土壤氮素和氮磷比呈负向响应,而磷重吸收效率与氮磷比呈显著正相关,与土壤速效磷呈负相关。表明土壤养分有效性的变化负向驱动养分重吸收效率。由于该生境中刺槐林的固氮效应及磷限制,叶片养分重吸收策略对土壤氮磷比响应强烈。     

增温对若尔盖高寒草本沼泽木里薹草氮磷重吸收的影响

养分重吸收是植物重要的营养保存机制和养分循环的重要组成部分,温度变化会影响植物养分吸收。为了探讨若尔盖高原沼泽湿地植物木里薹(Carex muliensis)草养分重吸收特征对气候变暖的响应,本研究通过野外模拟增温实验,测定木里薹草成熟叶片和衰老叶片的氮(N)、磷(P)含量并分析其重吸收效率差异。结果表明：木里薹草叶片N、P含量均值分别为11.44和1.19 mg·g^-1,N重吸收效率(NRE)、P重吸收效率(PRE)均值分别为61.8%和69.0%,增温显著降低了成熟叶片氮含量,显著提高了衰老叶片磷含量(P<0.01),对成熟叶片磷含量和衰老叶片氮含量没有显著影响;增温显著降低了成熟叶片和衰老叶片N∶P(P<0.01)及NRE(P<0.05)和PRE(P<0.01);木里薹草叶片氮磷重吸收效率与成熟叶片氮磷含量呈显著正相关,与衰老叶片氮磷含量呈显著负相关;木里薹草生长受P限制,而增温可能导致限制情况发生变化,木里薹草叶片养分重吸收还可能受到化学计量调控。研究结果将有助于了解和预测若尔盖高原养分循环对未来气候变化的响应,并为气候变暖下高寒草...     

亚热带森林菌根植物根系真菌群落结构对氮磷添加的响应

植物根系真菌对维系植物的营养吸收和健康具有重要作用。本研究分析了不同菌根类型植物根系中真菌的群落结构对外源氮(N)、磷(P)、氮和磷(NP)输入的响应。试验采集了无添加(对照)和N、P、NP添加处理下亚热带森林3种菌根类型(丛枝菌根、外生菌根、欧石楠菌根)9种植物的根系，运用高通量测序技术检测根系中真菌的多样性和群落组成。结果表明：9种植物根系真菌群落均主要由担子菌门和子囊菌门组成；P添加下子囊菌门的相对多度显著低于对照，而担子菌门的相对多度显著高于对照。欧石楠菌根植物根系中子囊菌门的相对多度显著高于丛枝菌根和外生菌根植物，而其担子菌门的相对多度显著低于丛枝菌根和外生菌根植物。与对照相比，P添加显著降低了植物根系中真菌的α多样性，改变了不同菌根类型植物根系中真菌的群落组成，而N添加和菌根类型的影响不明显。与对照和N添加相比，NP添加使全部植物根系中真菌群落变异程度更大，即群落整体更加趋异，而外生菌根植物根系中的真菌群落比丛枝菌根植物根系中的真菌群落变异更小，即群落更趋同。综上，P养分是影响亚热带森林土壤中树木根系真菌群落结构的关键因素。本研究有助于提升对全球环境变化下亚热带地区植物根系...     

罗浮栲和米槠细根形态功能性状对短期氮添加的可塑性响应

氮沉降会影响细根的形态功能性状,进而影响细根对养分的吸收,导致陆地生态系统养分循环发生变化.为了解氮沉降对细根形态功能性状的影响,利用根袋法进行原位试验,研究中亚热带常绿阔叶林外生菌根树种罗浮栲和米槠细根形态对短期氮添加的可塑性响应.结果表明: 低序级根(1~3序级)的比根长和比表面积对氮添加的可塑性响应高于高序级根(4序级),细根组织密度对氮添加的可塑性响应从1序级到4序级逐渐加强,而各序级细根直径对氮添加则无显著的可塑性响应;低序级细根比根长、比表面积的可塑性响应与高序级细根组织密度的可塑性响应之间存在一定的协同变化.罗浮栲和米槠细根的比根长、比表面积、组织密度对氮添加表现出相反方向的可塑性响应,表明施氮后不同外生菌根树种采取了不同的养分觅食策略:施氮后罗浮栲在养分获取上采取的是增加比根长、比表面积和根长增殖速率的资源快速获取策略,而米槠则采取了增大细根组织密度的相对保守的资源获取策略.     

Coloration and phenology manifest nutrient variability in senesced leaves of 46 temperate deciduous woody species

颜色和物候表明46种温带落叶木本植物衰老叶片的养分变异 不同共生植物的叶片养分含量差异显著,反映了不同的叶片养分利用策略。然而,衰老叶片养分的种间变异及其驱动因素尚不清楚。本研究旨在探讨衰老叶片养分的种间变异及其驱动因素。我们在中国东北的帽儿山森林生态系统研究站测定了46种共存温带落叶木本植物新鲜落叶的碳、氮、磷浓度。 采用随机森林模型量化10个生物因素(菌根类型、固氮类型、生长形态、耐阴性、叶片质地、变色程度、变色类型、叶片变色峰期、落叶峰期和落叶末期)的相对重要性。研究结果表明,落叶氮浓度种间变化为4倍,磷浓度变化达9倍。较高的氮和磷平均浓度(15.38和1.24 mg g⁻¹)表明该森林氮和磷限制较弱。功能群仅对特定养分及其比值有显著影响。磷浓度、氮磷比与外生菌根树种的落叶高峰日和落叶结束日呈负相关。颜色鲜艳的叶片(红色＞棕色＞黄色＞黄绿色＞绿色)倾向于比绿色叶片氮和磷浓度更低而碳氮比和碳磷比较高。随机森林模型表明,秋季叶变色和落叶物候贡献了80%的种间变异解释量。这些结果增加了我们对温带森林木本植物营养策略之衰老叶片养分变异性的理解。     

Resorption efficiency of leaf nutrients in woody plants on Mt. Dongling of Beijing,North China

Aims To explore resorption efficiency of nitrogen (NRE) and phosphorus (PRE) of woody plants in relation to soil nutrient availability, climate and evolutionary history, in North China.Methods We measured concentrations of nitrogen ([N]) and phosphorus ([P]) in both full expanded mature green and senescent leaves of the same individuals for 88 woody species from 10 sites of Mt. Dongling, Beijing, China. We built a phylogenetic tree for all these species and compared NRE and PRE among life forms (trees, shrubs and woody lianas) and between functional groups (N-fixers and non-N-fixers). We then explored patterns of NRE and PRE along gradients of mean annual temperature (MAT), soil inorganic N and available P, and phylogeny using a general linear model.Important findings Mass-based NRE (NRE m) and PRE (PRE m) averaged 57.4 and 61.4%, respectively, with no significant difference among life forms or functional groups. Neither NRE m nor PRE m exhibited significant phylogenetic signals, indicating that NRE m and PRE m were not phylogenetically conserved. NRE m was not related to [N] in green leaves; PRE m was positively correlated with [P] in green leaves; however, this relationship disappeared for different groups. NRE m decreased with [N] in senescent leaves, PRE m decreased with [P] in senescent leaves, for all species combined and for trees and shrubs. NRE m decreased with soil inorganic N for all species and for shrubs; PRE m did not exhibit a significant trend with soil available P for all species or for different plant groups. Neither NRE m nor PRE m was significantly related to MAT for overall species and for species of different groups.     

江西阳际峰30种阔叶树叶片氮磷含量及再吸收效率

养分再吸收是植物养分利用的重要策略,体现了植物对养分留存、利用和适应环境的能力。为研究亚热带不同生活型(常绿与落叶)阔叶树养分含量与养分再吸收的关系,以江西阳际峰国家级自然保护区内30种阔叶树为研究对象,测定成熟和衰老叶片氮(N)和磷(P)含量,分析常绿和落叶树种叶片N和P含量及其再吸收效率差异,揭示阔叶树种叶片养分再吸收效率对植物生活型的响应。结果表明: 落叶树种成熟叶片N和P含量显著高于常绿树种,衰老叶片P含量显著高于常绿树种,而两者衰老叶N含量差异不显著;30种阔叶林木叶片的氮再吸收效率(NRE)与磷再吸收效率(PRE)平均值分别为49.6%和50.9%,两种生活型树种间叶片的NRE与PRE无显著差异;落叶和常绿树种叶片的NRE均与衰老叶N含量呈显著负相关,PRE则与衰老叶P含量呈显著负相关,且这种关系在不同生活型之间差异不显著;总物种的PRE-NRE异速生长指数为1.18。江西阳际峰30种不同生活型阔叶树的养分再吸收效率会影响衰老叶片的养分状况,且相较于N,植物偏好从衰老叶中再吸收P。     

广西猫儿山不同海拔常绿和落叶树种的营养再吸收模式

土壤养分供给性大小是否影响植物氮和磷再吸收效率仍存在争议。调查了广西猫儿山不同海拔常绿和落叶树种成熟和衰老叶片的氮和磷含量,探讨营养再吸收是否受到叶片习性和海拔的影响。所有树种氮和磷再吸收效率的平均值分别为56.5%和52.1%。常绿树种比落叶树种有显著较高的氮再吸收效率(P0.001)和磷再吸收效率(P0.01),这与前者有较低的衰老叶片氮和磷含量密切相关。随着海拔的上升,氮再吸收效率显著下降(P0.01),磷再吸收效率显著提高(P0.05)。氮再吸收效率与土壤氮:磷比(r=-0.41,P0.05)和成熟叶片氮:磷比(r=-0.37,P0.05)负相关,磷再吸收效率与土壤氮:磷比(r=0.44,P0.05)和成熟叶片氮:磷比(r=0.47,P0.01)正相关,表明了树种对低海拔氮限制的适应逐渐转变为对高海拔磷限制的适应。此外,氮再吸收效率与年均温正相关(r=0.43,P0.05)而磷再吸收效率与年均温负相关(r=-0.45,P0.01),这表明气温也是调节树木营养再吸收格局的重要影响因素。不同海拔树种氮和磷再吸收模式的差异可能是引起广西猫儿山常绿树种沿海拔形成双峰分布的原因之一。     

Resorption proficiency and efficiency of leaf nutrients in woody plants in eastern China

Aims (i) To explore variations in nutrient resorption of woody plants and their relationship with nutrient limitation and (ii) to identify the factors that control these variations in forests of eastern China.Methods We measured nitrogen (N) and phosphorus (P) concentrations in both green and senesced leaves of 172 woody species at 10 forest sites across eastern China. We compared the nutrient resorption proficiency (NuRP) and efficiency (NuRE) of N and P in plant leaves for different functional groups; we further investigated the latitudinal and altitudinal variations in NuRP and NuRE and the impacts of climate, soil and plant types on leaf nutrient resorptions.Important findings On average, the leaf N resorption proficiency (NRP) and P resorption proficiency (PRP) of woody plants in eastern China were 11.1mg g ? 1 and 0.65 mg g ? 1, respectively; and the corresponding N resorption efficiency (NRE) and P resorption efficiency (PRE) were 49.1% and 51.0%, respectively. Angiosperms have higher NRP (are less proficient) values and lower NRE and PRE values than gymnosperms, but there are no significant differences in NRP, PRP and PRE values between species with different leaf habits (evergreen vs. deciduous angiosperms). Trees have higher NRE and PRE than shrubs. Significant geographical patterns of plant nutrient resorption exist in forests of eastern China. In general, NRP and PRE decrease and PRP and NRE increase with increasing latitude/altitude for all woody species and for the different plant groups. Plant functional groups show more controls than environmental factors (climate and soil) on the N resorption traits (NRP and NRE), while site-related variables present more controls than plant types on PRP and PRE. NRP increases and PRP and NRE decrease significantly with increasing temperature and precipitation for the overall plants and for most groups, except that significant PRE–climate relationship holds for only evergreen angiosperms. Leaf nutrient resorption did not show consistent responses in relation to soil total N and P stoichiometry, probably because the resorption process is regulated by the relative costs of drawing nutrients from soil versus from senescing leaves. These results support our hypothesis that plants growing in P-limited habitats (low latitudes/altitudes or areas with high precipitation/temperature) should have lower PRP and higher PRE, compared with their counterparts in relatively N-limited places (high latitudes/altitudes or areas with low precipitation/temperature). Our findings can improve the understanding of variations in N and P resorption and their responses to global change, and thus facilitate to incorporate these nutrient resorption processes into future biogeochemical models.     

N limitation increases along a temperate forest succession: evidences from leaf stoichiometry and nutrient resorption

温带森林演替加剧了氮限制：来自叶片化学计量和养分重吸收的证据 森林生产力和碳汇功能在很大程度上取决于土壤氮和磷的有效性。然而,迄今为止,养分限制随森林演替的时间变化仍存在争议。叶片化学计量和养分重吸收是预测植物生长养分限制的重要指标。基于此,本研究测定了温带森林4个演替阶段所有木本植物叶片和凋落叶中氮和磷的含量,并分析了演替过程中非生物因子和生物因子如何影响叶片化学计量和养分重吸收。研究结果表明,在个体尺度上,叶片氮磷含量在演替末期显著增加,而叶片氮磷比无显著变化;氮的重吸收效率随演替显著增加,然而磷的重吸收效率先增加后减少;氮重吸收效率与磷重吸收效率的比值仅在演替末期显著增加。此外,植物氮素循环对土壤养分的响应比磷素循环更弱。在群落尺度上,叶片氮磷含量随森林演替呈现先降低后升高的趋势,主要受香农-维纳多样性指数和物种丰富度的影响;叶片氮磷比随演替而显著变化,主要由胸径的群落加权平均值决定;氮的重吸收效率增加,主要受物种丰富度和胸径的影响,而磷的重吸收效率相对稳定。因此,氮重吸收效率与磷重吸收效率的比值显著增加,表明随着温带森林演替,氮限制加剧。这些结果可能反映了较高生物多样性群落中物种间对有限资源的激烈竞争,强调了生物因子在驱动森林生态系统养分循环中的重要性,为中国温带和北方森林可持续经营的施肥管理提供了参考。     

Stoichiometric and nutrient resorption characteristics of dominant tree species in subtropical Chinese forests

This study investigated seasonal patterns in stoichiometric ratios, nutrient resorption characteristics, and nutrient use strategies of dominant tree species at three successional stages in subtropical China, which have not been fully understood. Fresh leaf and leaf litterfall samples were collected in growing and nongrowing seasons for determining the concentrations of carbon (C), nitrogen (N), and phosphorus (P). Then, stoichiometric ratios (i.e., C:N, C:P, N:P, and C:N:P) and resorption parameters were calculated. Our results found that there was no consistent variation in leaf C:N and C:P ratios among different species. However, leaf N:P ratios in late‐successional species became significantly higher, indicating that P limitation increases during successional development. Due to the P limitation in this study area, P resorption efficiency and proficiency were higher than corresponding N resorption parameters. Dominant tree species at early‐successional stage adopted “conservative consumption” nutrient use strategy, whereas the species at late‐successional stage inclined to adopt “resource spending” strategy.     

氮添加对黄河三角洲滨海湿地芦苇养分再吸收效率的影响

大气氮沉降增加能改变土壤养分可利用性,影响滨海湿地植物的养分再吸收。目前研究多关注氮沉降量对养分再吸收过程的影响,且研究集中于叶片,鲜有研究区分不同形态氮素对植物不同器官养分再吸收过程的影响。通过两年的野外控制实验,研究硝态氮、铵态氮添加对黄河三角洲滨海湿地芦苇(Phragmites australis)叶、茎养分再吸收效率的影响。结果表明：两类氮添加均显著增加叶、茎的氮、磷含量(P<0.001),增幅达32.74%—43.22%(氮)、30.91%—36.51%(磷)。叶片氮的再吸收效率为54.14%—67.66%,茎氮的再吸收效率为50.60%—62.85%。叶片磷的再吸收效率为56.80%—70.38%,茎磷的再吸收效率为77.43%—84.95%。两类氮添加均显著降低氮、磷的再吸收效率(P<0.001),但两类氮添加处理下的养分再吸收效率无差异。叶、茎氮的再吸收效率无差异,但茎磷的再吸收效率明显高于叶(P<0.01)。总之,氮添加降低芦苇对氮、磷的再吸收效率,且茎对养分的再吸收也具有不可忽略的贡献。     

Effects of artificial grassland establishment on soil nutrients and carbon properties in a black-soil-type degraded grassland

Despite a growing knowledge of nutrient limitation for mangrove species and how mangroves adapt to low nutrients, there is scant information about the relative importance of N:P ratio and leaf phenolics variability in determining nutrient conservation. In this study, we evaluated possible nutrient conservation strategies of a mangrove Rhizophora stylosa under nutrient limitation. 1. The leaf nutrient concentrations of R. stylosa changed with season, with the highest N concentration in winter and the highest P concentration in spring for both mature and senescent leaves. Leaf N and P concentrations decreased significantly during leaf senescence. Based on N:P ratios R. stylosa forest was N-limited. Accordingly, the nitrogen resorption efficiency (NRE) was significantly higher than phosphorus resorption efficiency (PRE) for the R. stylosa leaves during leaf senescence. The NRE and PRE both reached the highest in the autumn. Average N and P concentrations in the senescent leaves were 0.15% and 0.06% for R. stylosa, respectively, indicating a complete resorption of N and an incomplete resorption of P. There was a significant negative correlation between nitrogen resorption proficiency (NRP) and NRE, meanwhile phosphorus resorption proficiency (PRP) and PRE correlation was also highly significantly. 2. R. stylosa leaves contained relatively high tannin level. Total phenolics, extractable condensed tannins and total condensed tannins contents increased during leaf senescence, and changed between seasons. The lowest concentrations of total phenolics, extractable condensed tannins and total condensed tannins occurred in summer, total phenolics concentrations were inversely related to nitrogen or phosphorus concentrations. 3. Our results confirmed that resorption efficiency during leaf senescence depends on the type of nutrient limitation, and NRE was much higher than PRE under N-limited conditions. R. stylosa forest developed several nutrient conservation strategies in the intertidal coastline surroundings, including high nitrogen resorption efficiency, low nutrient losses and high tannins level.     

Inter‐specific Variation in Foliar Nutrients and Resorption of Nine Canopy‐tree Species in a Secondary Neotropical Rain Forest

The influence of environmental gradients on the foliar nutrient economy of forests has been well documented; however, we have little understanding of what drives variability among individuals within a single forest stand, especially tropical forests. We evaluated inter‐ and intra‐specific variation in nutrient resorption, foliar nutrient concentrations and physical leaf traits of nine canopy tree species within a 1‐ha secondary tropical rain forest in northeastern Costa Rica. Both nitrogen (N) and phosphorus (P) resorption efficiency (RE) and proficiency of the nine tree species varied significantly among species, but not within. Both N and P RE were significantly negatively related to leaf specific strength. Green leaf N and P concentrations were strongly negatively related to leaf mass per area, and senesced leaf nutrient concentrations were significantly positively related to green leaf nutrient concentrations. This study reveals a strong influence of physical leaf traits on foliar nutrient and resorption traits of co‐occurring species in a secondary wet tropical forest stand.