内蒙古锡林河流域草原群落土壤呼吸的时空变异及其影响因子研究

　　土壤呼吸是全球碳循环中的一个重要环节，其对全球碳平衡的影响是近年来人们关注的焦点之一。探讨碳素的失汇(missing sink)问题，对陆地生态系统土壤呼吸的研究是必不可少的。环境因子与土壤呼吸之间的关系可以用于将土壤呼吸从“chamber”水平的测量放大到整个生态系统或更大尺度。而温度、水分和植被状况都是对土壤呼吸有重要影响的因子，随着全球气候的变化，这些因子也会发生相应的改变，在这种情况下，它们极有可能与土壤CO2排放之间形成正反馈。温带草原是主要的陆地生态系统类型之一，目前非常缺乏有关土壤呼吸的研究资料。因此，在2001年生长季，我们在内蒙古锡林河流域南部集水区设定了一条东西长约160km、南北宽约30km的样带，从中选择了11个不同的植物群落，采用碱液吸收法周期性地对这些群落的土壤呼吸速率进行同步测定，并对土壤呼吸的时空动态及其与温度、土壤水分和植被状况之间的关系进行了研究。现将主要研究结果概述如下： 　　①锡林河流域南部集水区的土壤呼吸表现出明显的季节变化和空间变异。温度是影响土壤呼吸季节变化的主要因子之一，指数模型能够较好地揭示各群落土壤呼吸对温度变化的响应，但低温时模型的拟合效果更好。各群落土壤呼吸的季节动态与温度变化不完全同步，表明温度并不是影响土壤呼吸的唯一因子 。 　　②土壤呼吸的温度敏感性在各群落之间存在着一定的差异。春小麦群落的Q10值高于草原群落，说明不同的土地利用方式会影响到土壤呼吸对温度变化的敏感程度。水分对土壤呼吸的温度敏感性有重要影响，秩相关分析的结果表明，土壤水分与Q10值之间存在着显著的正相关关系。此外，依据不同土壤层次的温度得出的Q10值各不相同，基于变化幅度大的浅层土壤温度和气温得出的Q10值较小，而根据变化幅度小的深层土壤温度得出的Q10值较大。 　　③水分对各群落的土壤呼吸也有较大影响，但其影响程度有一定的季节差异，生长旺季水分对土壤呼吸的影响显著高于其它季节。从各群落的具体情况来看，水分对土壤呼吸的影响明显受制于群落的水分供应状况。水分供应状况比较好的和水分变化幅度小的群落中，土壤呼吸与水分之间没有显著的函数关系，而水分相对欠缺的群落则存有显著的线性关系。消除温度的影响后，这种线性关系显著增强。土壤水分含量较低的芨芨草群落中，土壤呼吸与表层水分之间的关系也不明显，这与芨芨草根系分布较深，能够利用土壤中较深层次的水分有关。 　　④土壤呼吸季节变化与植被之间的关系与各群落内水分状况以及植被对水分的利用机制有关。所有群落土壤呼吸速率随着绿色活体生物量的增长有上升趋势，且在水分供应充足的群落和植被较为耐旱或能够利用深层土壤水的群落中，这二者之间呈显著或极显著的指数关系，其它群落中相关关系不够显著。由于植被立枯量大小反映了水热的综合状况，所以群落的土壤呼吸速率随立枯量的增长呈下降趋势，二者之间的关系也可以用指数方程来表示。 　　⑤土壤呼吸在锡林河流域南部的空间变异主要受水分和植被状况的影响。总体来看，土壤水分含量高、地上生物量（包括绿色活体生物量）大或地上净第一性生产力高的草地群落，其土壤呼吸速率也较高。基础呼吸速率对于改进土壤呼吸模型在时间和空间上的预测精度有重要意义。我们的研究结果表明，在平均温度低、水分状况好、地上和地下生物量大、地上净第一性生产力高的地方，基础土壤呼吸速率也相应较高。

锡林河流域温带草地植物VOC释放及其对草地生态系统碳循环的贡献

植物源挥发性有机碳化合物（Volitale organic compounds, VOC）是大气VOC的主要来源，与对流层大气质量、大气化学密切相关。鉴于温带草地的分布范围很广，草地植物VOC释放潜力某种程度上影响植物源VOC的总释放量。另外，植物源VOC也是光合作用固定碳素的损失方式之一，可能在特定区域或生态系统中具有重要意义。基于上述想法，本文设计了四个方面的实验作为研究内容：1） 温带草地物种水平VOC释放潜力、及其与植物功能群的关系？2) 沙地植物物种水平VOC释放潜力、及其与植物功能群的关系？3) 沙地植物-草地植物VOC释放潜力存在显著性差异吗？4) 温带典型草地和退化草地的VOC释放速率如何？在生态系统水平，植物源VOC对温带草地碳循环的贡献多大？ 在所测定的175种温带草地植物中，不同植物间异戊二烯和单萜释放潜力差异很大;除少数物种外，大多数植物的异戊二烯和单萜释放潜力都较低，尤其是典型草地的优势物种。在此基础上，作者探讨了分类学赋值方法对温带草地植被的可行性，并初步建立了锡林河流域温带草地植物的VOC释放目录（共277种植物）。另外，温带草地植物的异戊二烯和单萜释放潜力与植物功能群（植物生活型和水分功能群）具有一定的联系，尤其是植物生活型。总的来说，温带草原的优势生活型（物种），即多年生根茎禾草（或多年生丛生禾草），具有较低的异戊二烯和单萜释放潜力。各水分功能群间差异不显著，但中旱生植物、旱中生植物 (温带草原的优势功能群)，具有较低的异戊二烯、单萜释放潜力。因此，温带草原退化过程中，那些具有较高VOC释放潜力的植物，重要性将会增加。 沙地植物种类组成非常丰富，不同物种间的异戊二烯和单萜释放潜力变异也很大。另外，沙地植物的异戊二烯和单萜释放潜力与其功能群间关系较密切，不同植物生活型间差异显著;其中也以多年生根茎禾草、多年生从生禾草的释放潜力最低，而乔木的释放潜力相对最高;该结论基本与草地的研究结论一致。然而，沙地植物的异戊二烯和单萜释放潜力与其水分功能群的关系比较模糊，中生植物具有更高的释放潜力，湿生植物的释放潜力较小。 通过对比沙地植物和草地植物的释放潜力，发现沙地植物的异戊二烯和单萜释放潜力比草地植物高，且这种差异整体上显著。另外，这种显著性差异，在不同植物生活型、水分功能群间也同样存在。沙地植物比对应的草地植物具有更高的异戊二烯和单萜释放潜力，最可能的原因：沙地正午的温度明显比草地温度高，前者实测温度可超过 45 ℃,这种经常性、周期性高温，促使沙地植物采用与草地植物不同的适应策略，即沙地植物通过释放更多的异戊二烯或单萜来减少其可能遭的热胁迫或热伤害，这种长期适应策略，使沙地植物具有更高的萜类化合物释放潜力。 本文还调查了温带典型草地生态系统和退化草地生态系统的异戊二烯和单萜释放速率，结果表明典型草地的标准异戊二烯和单萜释放速率分别为0.50 μgC g-1 h-1和0.69 μgC g-1 h-1;退化草地的标准异戊二烯和单萜释放潜力分别为0.32 μgC g-1 h-1和1.59 μgC g-1 h-1。总的来说，温带草地的异戊二烯和单萜释放速率都比较低，尤其是典型草地。整个生长季，典型草地释放的异戊二烯和单萜分别为31.6 mgC•m-2　和 70.4 mgC•m-2;退化草地的异戊二烯和单萜释放量分别为20.8 mgC•m-2 和 168.8 mgC•m-2。退化草地萜类化合物总释放速率远高于典型草地，尤其是单萜释放能力。过度放牧引起的草地退化，通过改变植被种类组成，对温带草地的异戊二烯和单萜释放速率产生显著影响;总体而言，温带草地退化将会使草地释放更多萜类化合物。 在温带草地生态系统中，Clost as VOC相对其NPP而言很小，在环境PAR和温度高时,它的贡献率相对较大;Clost as VOC约占典型草地生态系统NEP的5.32 %，退化草地生态系统NEP的0.23 %。植物源VOC释放所损失的碳素相对草地生态系统NPP而言几乎可以忽略不计;但是，相对其NEP，Clost as VOC还是具有一定的相关性。虽然，草地生态系统Clost as VOC对NPP或NEP的贡献率较小，但考虑到全球尺度植物源VOC的巨大释放速率，它在碳循环中的贡献率仍然不容忽视;在某些特殊的生态系统中，仍可能扮演重要角色。

内蒙古温带典型草原土壤净氮 矿化作用

作为陆地植物生长和生态系统初级生产力的主要限制因子之一，土壤氮素矿化和可利用性对全球变化的响应决定着未来陆地生态系统的碳储量并对全球碳循环产生长远的深刻影响。众所周知，土壤中大部分氮以有机态存在，有机态氮必须经过转化成为无机态氮才能被植物吸收利用。土壤氮素矿化是将有机态氮转化为无机态氮的生物化学过程，决定着土壤中氮素的可利用性。理解氮素在草原生态系统中的转化（包括矿化，硝化作用）过程，有助于我们充分认识草地退化机理，为草地恢复和重建提供理论依据。在中国科学院内蒙古草原生态系统定位站的典型羊草草原区，选取了1979年围封（没有割草和放牧利用）、1999年围封（没有割草和放牧利用）和长期自由放牧三种不同土地利用方式的羊草草地，分别代表人为干扰强度从小到大的梯度，利用室内和野外原状土培育实验两种方法，研究了人为干扰强度和环境因子（温度和水分）对土壤无机氮转化的影响。 室内实验：设置不同的温度梯度和水分梯度，在不同的时间段内在室内对原状土柱进行培养，观察这些因子对土壤净氮矿化的影响。结果表明：（1）土壤无机氮库（NH4+-N和NO3−-N）在不同土地利用变化情况下具有显著差异。但是，经过室内培养，净氮矿化速率和硝化速率之间差异不显著，只有铵化速率表现出显著差异。（2）温度对铵化、硝化和矿化速率有显著影响。但是当培养温度低于5℃时，无论培养时间多长净氮矿化的累积和净氮矿化速率之间差异皆不显著。温度高于15℃的三个温度之间净氮矿化的累积和净氮矿化速率差异极显著。（3）土壤湿度对净硝化和净矿化速率有显著影响，但是对铵化速率没有显著影响，表明当土壤水分限制硝化细菌的活性时，硝化速率对于水分的增加显得更加敏感。（4）培养时间对铵化、硝化和矿化量的积累有极显著影响。随着培养时间的延长铵态氮积累的量较少，但是硝态氮的量随着培养时间延长积累的量很多。（5）我们的研究表明温度、湿度和培养时间之间对净氮矿化的影响存在极显著的互作效应。 野外试验：在三种不同土地利用历史的草地，利用顶盖埋管原位培育法测定土壤的无机氮库、净硝化、铵化和净氮矿化的季节动态。2004年5月开始，每隔30天一次，到11月结束。结果表明：（1）三种不同土地利用方式的草地生态系统土壤中的NO3－-N和NH4+-N含量都表现出明显的季节变化趋势，自由放牧样地与围封25年样地季节趋势基本一致，围封5年样地各月间NO3－-N和NH4+-N含量的差异都达到显著水平。（2）三种不同土地利用方式的草地无机氮的季节动态两两比较发现，NO3－-N浓度在6、7和9月差异极显著;而在8月三个样地之间差异不显著;NH4+-N浓度在5月、7月和8月三个样地之间都达到显著水平，在6月、9月和10月情况有所不同，6月自由放牧地与围封25年间差异不显著，而这两个样地与围封5年的样地差异达显著水平，9月围封25年和围封5年两个样地之间差异不显著，而与自由放牧样地之间差异显著;10月情况又有变化，是自由放牧样地与围封5年的样地之间差异不显著，与围封25年样地之间差异达显著水平;（3）土壤无机氮库和氮素的矿化/硝化速率都存在明显且比较一致的季节动态，但是在个别月份也有较大差异，总的趋势是围封25年的样地硝化速率高于围封5年和自由放牧样地。上述结果表明：在温带典型草原，土地利用方式对土壤氮素库和矿化有着重要影响;各样地之间土壤氮素库和矿化速率的差异可能是由于土壤温度、水分、无机氮库和土壤基质的改变而引起的。

改变土壤碳输入对我国北方温带草原土壤碳、氮库和碳、氮循环的影响

土壤是陆地生态系统最重要的碳库和氮库，在陆地生态系统碳氮循环过程中起着举足轻重的作用。植物光合作用产物向地下的分配(根系分泌、细胞凋亡和根系分解)和地上凋落物分解是土壤碳两个最主要的来源，关于这两种碳输入途径和量如何影响陆地生态系统土壤碳氮库和循环的有限理解，限制了我们预测陆地生态系统碳循环对于全球气候变化的响应和反馈。本实验以我国北方广泛分布的温带典型草原生态系统为研究对象，通过添加和去除地面凋落物以及割草改变植物对土壤的碳输入途径和量，观察温带草原土壤有机碳库和氮库各个组分(微生物生物量碳氮、活性碳、可溶性碳、轻重组碳、总有机碳、总有机氮)及碳氮循环(微生物呼吸、土壤呼吸、生态系统气体交换、净氮矿化)的变化，来区分和量化凋落物分解和根系分泌、分解这两种碳输入方式对草原土壤碳库和循环的相对贡献和影响，有助于揭示在全球变化背景下温带典型草原生态系统碳库和碳循环的响应机制，并为预测其未来变化动态以及对全球气候变化的反馈提供参数估计和模型校正和验证。 我们对各处理样方土壤温度、土壤水分以及土壤呼吸进行了一个生长季的测定，结果表明，添加和去除凋落物都使得土壤呼吸与对照相比都有所增加，但是都没有达到显著水平;而无论地上凋落物的量添加、去除还是不变，植物光合作用对土壤呼吸的影响则比较显著，去除植物显著地降低了土壤呼吸。这说明处理一年后，植物对土壤呼吸起主导作用，而凋落物处理对土壤呼吸的影响还需要更长时间的观察。此外，生物量的结果也表明了地上凋落物的改变会影响到植物生长，从而影响到植物对土壤的碳输入，但是都没有达到显著水平，仍然需要我们进行更长时间的处理和观察。 通过在生长季前期、中期、后期进行了三次氮矿化的测定，以及在生长季后期进行了一次微生物生物量测定，我们得出以下结论：添加凋落物加快了净氮矿化速率，但是没有达到显著影响水平;同样，去除凋落物降低了净氮矿化速率，也没有达到显著水平。此外，添加凋落物处理微生物生物量碳、氮都有所增加，与对照相比没有显著差异，但是显著高于另外几个处理。 通过本研究，我们对于根系分泌和地上凋落物分解这两种碳氮输入途径对我国温带草原碳氮循环的影响有了初步的了解。与森林生态系统相比，由于草原生态系统土壤碳氮含量低，土壤碳库和转化对碳输入途径和量的变化更为敏感，在处理第一年，土壤微生物量和土壤呼吸都发生了显著的变化。然而，由于改变土壤碳库是长时间日积月累的结果，草原生态系统某些指标对碳输入途径和量的变化的响应还需要较长时间才能显示出来，例如添加和去除凋落物虽然使得土壤呼吸值有所增加，但是还都没有达到显著的水平;另外碳输入途径和量的变化对草原生态系统氮矿化还没有产生显著的影响。这些都需要更长时间的处理和观测来验证。

土壤有机物添加对温带半干旱草地地下生物过程的影响

大气CO2 浓度和降水量增加有可能大幅提高中国北方部分草地生态系统净初级生产力，进而导致向土壤中输送的有机物相应增加。本研究以位于内蒙古自治区东乌珠穆沁旗内的半干旱草地生态系统为研究对象，通过向10−20 cm 土层添加不同质量和数量的植物凋枯物碎屑模拟有机物输入增加和喷灌模拟降水量提升，同时测定土壤微生物群落动态和植物生长指标，探讨在增加有机物输入和土壤水分的情况下土壤生物过程的变化及其对土壤碳排放和贮存的反馈作用。 研究结果表明，有机物添加可促进植物地上部分及根系的生长，并显著提高土壤中可溶性有机碳（能量）和氮（养分）的含量。土壤能量和养分水平的提高促进了土壤微生物的生长：在底物可利用性水平较高时，r−对策微生物（指具有生长迅速、C/N 值较低的微生物群组）在群落中占优势地位；随着底物水平的降低，土壤中K-对策微生物（指具有生长缓慢、C/N 值较高的微生物群组）在群落中逐渐占据优势地位。土壤微生物群落组成的改变进一步导致了微生物功能群代谢活性和特征的变化，具体表现为提高了有机物添加处理中土壤细菌群落的代谢潜能，并使细菌在群落水平上的生理剖面特征明显区别于未添加有机物的处理。 研究样地内土壤微生物主要受到底物中的能量（碳）限制，土壤活性有机质库作为可利用性较高的能量和养分的重要来源，对土壤微生物活性和土壤碳周转起着比水分因子更加重要的作用。土壤水分主要影响植物生长和根系活性，并增加了土壤微生物对底物响应的复杂性，但它对地下生物过程的作用程度以底物中能量和养分水平为前提。 利用稳定性13C 同位素示踪技术测量后发现，添加C4-植物凋枯物会加速C3 底物中碳的分解速度。结合有机物添加后土壤有机质库的变化，可以推测植物凋枯物（即能量物质）输入增加会导致土壤原生有机碳的正向激发效应。在此过程中，土壤微生物群落组成及功能群代谢活性的变化起着至关重要的作用。 不同光合途径（C3 和C4）的植物和同一植物不同器官组织（地上部分和根系）的凋枯物添加对地下生物过程，特别是土壤微生物群落代谢功能的影响是不同的。在添加C3-草本凋枯物的处理中，土壤细菌群落主要利用的碳源为氨基酸类化合物；而在添加C4-植物凋枯物的处理中，土壤细菌群落主要利用的碳源为羧酸类化合物。 本研究在野外自然条件下证明了在能量缺乏的中国北方草地生态系统中，土壤有机物输入增加不但不会提高土壤有机碳库的大小，而且可能导致土壤原生有机碳的激发效应。在利用土壤呼吸与环境因子（如温度）的关系进行模拟预测土壤碳排放时，需要考虑不同生态系统底物中的能量和养分水平，以及土壤微生物和植物根系等地下生物过程对底物水平的适应性。

Characterizing evapotranspiration over a meadow ecosystem on the Qinghai-Tibetan Plateau

To characterize evapotranspiration (ET) over grasslands on the Qinghai-Tibetan Plateau, we examined ET and its relevant environmental variables in a Kobresia meadow from 2002 to 2004 using the eddy covariance method. The annual precipitation changed greatly, with 554, 706, and 666 mm a(-1) for the three consecutive calendar years. The annual ET varied correspondingly to the annual precipitation with 341, 407, and 426 mm a(-1). The annual ET was, however, constant at about 60% of the annual precipitation. About 85% annual ET occurred during the growing season from May to September, and the averaged ET for this period was 1.90, 2.23, and 2.22 mm/d, respectively for the three consecutive years. The averaged ET was, however, very low (< 0.40 mm/d) during the nongrowing season from October to April. The annual canopy conductance (gc) and the Priestley-Taylor coefficient (a) showed the lowest values in the year with the lowest precipitation. This study first demonstrates that the alpine meadow ecosystem is characterized by a low ratio of annual ET to precipitation and that the interannual variation of ET is determined by annual precipitation.

Short-term variation of CO2 flux in relation to environmental controls in an alpine meadow on the Qinghai-Tibetan Plateau

[1] The alpine meadow ecosystem on the Qinghai-Tibetan Plateau may play a significant role in the regional carbon cycle. To assess the CO2 flux and its relationship to environmental controls in the ecosystem, eddy covariance of CO2, H2O, and energy fluxes was measured with an open-path system in an alpine meadow on the plateau at an elevation of 3,250 m. Net ecosystem CO2 influx (Fc) averaged 8.8 g m(-2) day(-1) during the period from August 9 to 31, 2001, with a maximum of 15.9 g m(-2) day(-1) and a minimum of 2.3 g m(-2) day(-1). Daytime Fc averaged 16.7 g m(-2) day(-1) and ranged from 10.4 g m(-2) day(-1) to 21.7 g m(-2) day(-1) during the study period. For the same photosynthetic photon flux density (PPFD), gross CO2 uptake (Gc) was significantly higher on cloudy days than on clear days. However, mean daily Gc was higher on clear days than on cloudy days. With high PPFD, Fc decreased as air temperature increased from 10degreesC to 23degreesC. The greater the difference between daytime and nighttime air temperatures, the more the sink was strengthened. Daytime average water use efficiency of the ecosystem (WUEe) was 8.7 mg (CO2)(g H2O)(-1); WUEe values ranged from 5.8 to 15.3 mg (CO2)(g H2O)(-1). WUEe increased with the decrease in vapor pressure deficit. Daily albedo averaged 0.20, ranging from 0.19 to 0.22 during the study period, and was negatively correlated with daily Fc. Our measurements provided some of the first evidence on CO2 exchange for a temperate alpine meadow ecosystem on the Qinghai-Tibetan Plateau, which is necessary for assessing the carbon budget and carbon cycle processes for temperate grassland ecosystems.

Compositional divergence and convergence in arbuscular mycorrhizal fungal communities

In spite of the controversy that they have generated, neutral models provide ecologists with powerful tools for creating dynamic predictions about beta-diversity in ecological communities. Ecologists can achieve an understanding of the assembly rules operating in nature by noting when and how these predictions are met or not met. This is particularly valuable for those groups of organisms that are challenging to study under natural conditions (e.g., bacteria and fungi). Here, we focused on arbuscular mycorrhizal fungal (AMF) communities and performed an extensive literature search that allowed us to synthesize the information in 19 data sets with the minimal requisites for creating a null hypothesis in terms of community dissimilarity expected under neutral dynamics. In order to achieve this task, we calculated the first estimates of neutral parameters for several AMF communities from different ecosystems. Communities were shown either to be consistent with neutrality or to diverge or converge with respect to the levels of compositional dissimilarity expected under neutrality. These data support the hypothesis that divergence occurs in systems where the effect of limited dispersal is overwhelmed by anthropogenic disturbance or extreme biological and environmental heterogeneity, whereas communities converge when systems have the potential for niche divergence within a relatively homogeneous set of environmental conditions. Regarding the study cases that were consistent with neutrality, the sampling designs employed may have covered relatively homogeneous environments in which the effects of dispersal limitation overwhelmed minor differences among AMF taxa that would lead to environmental filtering. Using neutral models we showed for the first time for a soil microbial group the conditions under which different assembly processes may determine different patterns of beta-diversity. Our synthesis is an important step showing how the application of general ecological theories to a model microbial taxon has the potential to shed light on the assembly and ecological dynamics of communities.

Choosing and using diversity indices: Insights for ecological applications from the German Biodiversity Exploratories

Biodiversity, a multidimensional property of natural systems, is difficult to quantify partly because of the multitude of indices proposed for this purpose. Indices aim to describe general properties of communities that allow us to compare different regions, taxa, and trophic levels. Therefore, they are of fundamental importance for environmental monitoring and conservation, although there is no consensus about which indices are more appropriate and informative. We tested several common diversity indices in a range of simple to complex statistical analyses in order to determine whether some were better suited for certain analyses than others. We used data collected around the focal plant Plantago lanceolata on 60 temperate grassland plots embedded in an agricultural landscape to explore relationships between the common diversity indices of species richness (S), Shannon's diversity (H'), Simpson's diversity (D1), Simpson's dominance (D2), Simpson's evenness (E), and Berger–Parker dominance (BP). We calculated each of these indices for herbaceous plants, arbuscular mycorrhizal fungi, aboveground arthropods, belowground insect larvae, and P. lanceolata molecular and chemical diversity. Including these trait-based measures of diversity allowed us to test whether or not they behaved similarly to the better studied species diversity. We used path analysis to determine whether compound indices detected more relationships between diversities of different organisms and traits than more basic indices. In the path models, more paths were significant when using H', even though all models except that with E were equally reliable. This demonstrates that while common diversity indices may appear interchangeable in simple analyses, when considering complex interactions, the choice of index can profoundly alter the interpretation of results. Data mining in order to identify the index producing the most significant results should be avoided, but simultaneously considering analyses using multiple indices can provide greater insight into the interactions in a system.

Palaeovegetation in China during the late Quaternary: Biome reconstructions based on a global scheme of plant functional types

Two previous reconstructions of palaeovegetation across the whole of China were performed using a simple classification of plant functional types (PFTs). Now a more explicit, global PFT classification scheme has been developed, and a substantial number of additional pollen records have become available. Here we apply the global scheme of PFTs to a comprehensive set of pollen records available from China to test the applicability of the global scheme of PFTs in China, and to obtain a well-founded reconstruction of changing palaeovegetation patterns. A total of 806 pollen surface samples, 188 mid-Holocene (MH, 6000 14C yr BP) and 50 last glacial maximum (LGM, 18,000 14C yr BP) pollen records were used to reconstruct vegetation patterns in China, based on a new global classification system of PFTs and a standard numerical technique for biome assignment (biomization). The biome reconstruction based on pollen surface samples showed convincing agreement with present potential natural vegetation. Coherent patterns of change in biome distribution between MH, LGM and present are observed. In the MH, cold and cool-temperate evergreen needleleaf forests and mixed forests, temperate deciduous broadleaf forest, and warm-temperate evergreen broadleaf and mixed forest in eastern China were shifted northward by 200–500 km. Cold-deciduous forest in northeastern China was replaced by cold evergreen needleleaf forest while in central northern China, cold-deciduous forest was present at some sites now occupied by temperate grassland and desert. The forest–grassland boundary was 200–300 km west of its present position. Temperate xerophytic shrubland, temperate grassland and desert covered a large area on the Tibetan Plateau, but the area of tundra was reduced. Treeline was 300–500 m higher than present in Tibet. These changes imply generally warmer winters, longer growing seasons and more precipitation during the MH. Westward shifts of the forest–shrubland–grassland and grassland–desert boundaries imply greater moisture availability in the MH, consistent with a stronger summer monsoon. During the LGM, in contrast, cold-deciduous forest, cool-temperate evergreen needleleaf forest, cool mixed forests, warm-temperate evergreen broadleaf and mixed forest in eastern China were displaced to the south by 300–1000 km, while temperate deciduous broadleaf forest, pure warm-temperate evergreen forest, tropical semi-evergreen and evergreen broadleaf forests were restricted or absent from the mainland of southern China, implying colder winters than present. Strong shifts of temperate xerophytic shrubland, temperate grassland and desert to the south and east in northern and western China and on the Tibetan Plateau imply drier conditions than present.

Grazing response patterns indicate isolation of semi-natural European grasslands

Identifying drivers of species diversity is a major challenge in understanding and predicting the dynamics of species-rich semi-natural grasslands. In particular in temperate grasslands changes in land use and its consequences, i.e. increasing fragmentation, the on-going loss of habitat and the declining importance of regional processes such as seed dispersal by livestock, are considered key drivers of the diversity loss witnessed within the last decades. It is a largely unresolved question to what degree current temperate grassland communities already reflect a decline of regional processes such as longer distance seed dispersal. Answering this question is challenging since it requires both a mechanistic approach to community dynamics and a sufficient data basis that allows identifying general patterns. Here, we present results of a local individual- and trait-based community model that was initialized with plant functional types (PFTs) derived from an extensive empirical data set of species-rich grasslands within the `Biodiversity Exploratories' in Germany. Driving model processes included above- and belowground competition, dynamic resource allocation to shoots and roots, clonal growth, grazing, and local seed dispersal. To test for the impact of regional processes we also simulated seed input from a regional species pool. Model output, with and without regional seed input, was compared with empirical community response patterns along a grazing gradient. Simulated response patterns of changes in PFT richness, Shannon diversity, and biomass production matched observed grazing response patterns surprisingly well if only local processes were considered. Already low levels of additional regional seed input led to stronger deviations from empirical community pattern. While these findings cannot rule out that regional processes other than those considered in the modeling study potentially play a role in shaping the local grassland communities, our comparison indicates that European grasslands are largely isolated, i.e. local mechanisms explain observed community patterns to a large extent.

Choosing and using diversity indices: insights for ecological applications from the German Biodiversity Exploratories

Biodiversity, a multidimensional property of natural systems, is difficult to quantify partly because of the multitude of indices proposed for this purpose. Indices aim to describe general properties of communities that allow us to compare different regions, taxa, and trophic levels. Therefore, they are of fundamental importance for environmental monitoring and conservation, although there is no consensus about which indices are more appropriate and informative. We tested several common diversity indices in a range of simple to complex statistical analyses in order to determine whether some were better suited for certain analyses than others. We used data collected around the focal plant Plantago lanceolata on 60 temperate grassland plots embedded in an agricultural landscape to explore relationships between the common diversity indices of species richness (S), Shannon's diversity (H'), Simpson's diversity (D-1), Simpson's dominance (D-2), Simpson's evenness (E), and Berger-Parker dominance (BP). We calculated each of these indices for herbaceous plants, arbuscular mycorrhizal fungi, aboveground arthropods, belowground insect larvae, and P.lanceolata molecular and chemical diversity. Including these trait-based measures of diversity allowed us to test whether or not they behaved similarly to the better studied species diversity. We used path analysis to determine whether compound indices detected more relationships between diversities of different organisms and traits than more basic indices. In the path models, more paths were significant when using H', even though all models except that with E were equally reliable. This demonstrates that while common diversity indices may appear interchangeable in simple analyses, when considering complex interactions, the choice of index can profoundly alter the interpretation of results. Data mining in order to identify the index producing the most significant results should be avoided, but simultaneously considering analyses using multiple indices can provide greater insight into the interactions in a system.

Collection of data on particulate and dissolved soil phosphorus in the Jena Experiment (Main Experiment, time series since 2002)

This data set contains two time series of measurements of dissolved phosphorus (organic, inorganic and total with a biweekly resolution) and dissolved inorganic phosphorus with a seasonal resolution. In addition, data on phosphorus from soil samples measured in 2007 and fractionated by different acid-extrations (Hedley fractions) are provided. All data measured at the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained by bi-annual weeding and mowing. 1. Dissolved phosphorus in soil solution: Suction plates installed on the field site in 10, 20, 30 and 60 cm depth were used to sample soil pore water. Cumulatively extracted soil solution was collected every two weeks from October 2002 to May 2006. The biweekly samples from 2002, 2003 and 2004 were analyzed for dissolved organic phosphorus (DOP), dissolved inorganic phosphorus (PO4P) and dissolved total phosphorus (TDP) by Continuous Flow Analyzer (CFA SAN ++, SKALAR [Breda, The Netherlands]). 2. Seasonal values of dissolved inorganic phosphorus in soil solution were calculated as volume-weighted mean values of the biweekly measurements (spring = March to May, summer = June to August, fall = September to November, winter = December to February). 3. Phosphorus fractions in soil: Five independent soil samples per plot were taken in a depth of 0-15 cm using a soil corer with an inner diameter of 1 cm. The five samples per plot were combined to one composite sample per plot. A four-step sequential P fractionation (Hedley fractions) was applied and concentrations of P fractions in soil were measured photometrically (molybdenum blue-reactive P) with a Continuous Flow Analyzer (Bran&Luebbe, Germany).

Mineral nitrogen from KCl extractions of soil from the Jena Experiment (Main Experiment up to 30cm depth, year 2002)

As an estimate of plant-available N, this data set contains measurements of inorganic nitrogen (NO3-N and NH4-N, the sum of which is termed mineral N or Nmin) determined by extraction with 1 M KCl solution of soil samples from the main experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained by bi-annual weeding and mowing. Soil sampling and analysis: Five soil cores (diameter 0.01 m) were taken at a depth of 0 to 0.15 m and 0.15 to 0.3 m of the mineral soil from each of the experimental plots in September 2002. Samples of the soil cores per plot were pooled during each sampling campaign. NO3-N and NH4-N concentrations were determined by extraction of soil samples with 1 M KCl solution and were measured in the soil extract with a Continuous Flow Analyzer (CFA, Skalar, Breda, Netherlands).