地球科学进展  2018 , 33 (4): 343-349 https://doi.org/10.11867/j.issn.1001-8166.2018.04.0343

科技重大计划进展

陆地系统中的露石及其生态作用

沈有信1, 赵志猛12, 毕胜春3, 赵高卷12, 刘娟12

1.中国科学院西双版纳热带植物园,云南 勐仑 666303
2.中国科学院大学,北京 100049
3. 石林花木有限责任公司,云南 石林 652208

Rock Outcrop and Its Ecological Function in Terrestrial Ecosystem

Shen Youxin1, Zhao Zhimeng12, Bi Shengchun3, Zhao Gaojuan12, Liu Juan12

1.Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun Yunnan 666303,China
2.University of Chinese Academy of Sciences, Beijing 100049,China
3.Shilin Flowers and Trees Co., Ltd., Shilin Yunnan 652208, China

中图分类号:  P951

文献标识码:  A

文章编号:  1001-8166(2018)04-0343-07

收稿日期: 2017-11-7

修回日期:  2018-03-20

网络出版日期:  2018-04-20

版权声明:  2018 地球科学进展 编辑部 

基金资助:  *国家重点研发计划项目“断陷盆地山地土壤流失过程与驱动机制”(编号:2016YFC0502503)国家自然科学基金项目“喀斯特生态系统裸露岩体集水径流及其生态水文效应研究”(编号:41671031)资助.

作者简介:

作者简介:沈有信(1966-),男,云南宜良人,研究员,主要从事喀斯特生态学与植被恢复研究工作.E-mail:yxshen@xtbg.ac.cn

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摘要

露石在陆地生态系统中十分常见,喀斯特生态系统尤为突出。露石在各种生态系统中产生积极或负面的影响,他们承接来自大气的降水(含穿透雨)、干湿沉降,殖居菌、藻、苔藓甚至高等维管束植物,可看成与邻近土壤斑块有鲜明对比的子生态系统。其承接和生产的水分和各种有机物质及无机物质输入邻近土壤斑块,影响水分、营养物质分配,进而对生长于土壤上的各种植物产生影响。定量测量露石承接、分配降水及其内含物的研究较少,评价其生态水文效应的研究更少,严重影响喀斯特生态系统演化机制的阐明与石漠化治理方略的制订。

关键词: 露石 ; 生物植居 ; 生态水文 ; 石漠化

Abstract

Rock Outcrop (ROC) is very common in terrestrial ecosystems, typically in karst. ROCs play both positive and negative effects in ecosystems. They may collect precipitation (including throughfall), wet/dry deposition, host bacteria, fungi, muss and lichen, and even vascular plants. Both plants and their growing matrixes on ROCs differ greatly from their nearby soil patches. Water and organic and inorganic materials received and produced on ROCs is easily redistributed to their nearby soil patches, put strong influence on water and elements process in soil patches, and thus, affect the plants growing on soil surface. However, quantitative study on water and materials received and produced is scarce, nor on the eco-hydrology effect, thus, blocking the explanation of karst ecosystem succession, block the strategy formulation on countermeasures of karst desertification.

Keywords: Rock outcrop ; Biological inhabitatione ; Eco-hydrology ; Rocky desertification.

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沈有信, 赵志猛, 毕胜春, 赵高卷, 刘娟. 陆地系统中的露石及其生态作用[J]. 地球科学进展, 2018, 33(4): 343-349 https://doi.org/10.11867/j.issn.1001-8166.2018.04.0343

Shen Youxin, Zhao Zhimeng, Bi Shengchun, Zhao Gaojuan, Liu Juan. Rock Outcrop and Its Ecological Function in Terrestrial Ecosystem[J]. Advances in Earth Science, 2018, 33(4): 343-349 https://doi.org/10.11867/j.issn.1001-8166.2018.04.0343

从地球的南北两极到赤道,从海拔高达几千米的冰川过渡带到接近海岸线,各种陆地生态系统中的裸露岩石个体十分常见。他们或是与地下基岩连接的尚未风化的基岩残余,或是被各种外营力搬运而远离原来基岩的碎石,其外形、大小、物理与化学性质各异,我们将其统称为“露石(rock outcrop)”。对于陆地生态系统的生物而言,无论这些裸露个体是否与基岩相连,其均会产生积极或负面的影响。

农业或林业上很早就关注到了土壤表面或是土体中的碎石(rock fragments)[1],他们是岩石到土壤的风化残余或是各种外营力搬运后的堆积物,与地下基岩无联系,个体小[2]。在干旱区,为了保水可在地表覆盖碎石[3]。中国热带、亚热带的碳酸盐经水力作用,形成了千姿百态的喀斯特地貌[46] 。此类地貌的自然林、人工林和石漠化迹地上,地表常见裸露的岩体(如石牙、溶峰和散落碎块,图1),这些独立或相连的不同形态、来源、大小、方位的裸露岩体或包围土壤斑块、或被土壤斑块包围(图2),共同组合出丰富而多样的喀斯特生态系统的植物生长基质。本文将更多关注此类露石,它们是生态系统中的自然产物,但长期以来未受到关注。

1 露石在陆地生态系统中的分布

沙漠(desert)、苔原(tundra)等植物生长困难的生态系统中,地表的露石很容易被观测到。各种权威的数据中均有含有露石的地类面积数据,如联合国粮农组织(FAO)发布的“Harmonized World Soil Database”(http:∥www.fao.org/geonetwork/srv/en/main.home)内的rock crust面积为1.2×106 km2,“Digital Soil Map of the World”(http:∥www.globalsoilmap.net/)给出rock crust的面积为0.4×106 km2。在全球性的土地利用类型分类及其分布图中,还有Barren等地类可能含有露石,如International Geo-sphere-Biosphere Programme 定义的Barren为:一年的任何时间段内植被覆盖均低于10%的、含有暴露的土壤、砂石、露石或是积雪的土地[7](遗憾的是,这些国际性的分类与分布图中,并未单独把rocks覆盖的面积区分出来,因此,尚无确切的统计数据显示含露石的陆地生态系统面积及其露石出露比例)。

图1   个体形状各异的不同岩溶生态系统露石

Fig.1   Karst rock outcrops with various sizes and shapes

全球的陆地生态系统中,喀斯特区域是被忽略了的含露石生态系统,其或许是全球面积最大的含露石生态系统。中国公布的涉及南方8个省(区、市)的石漠化和潜在石漠化(含裸露的岩体比例>30%)[8]面积是目前已知的最大喀斯特露石生态系统,其中2007年的面积分别为1.296×106和1.234×106 km2[9];2012年公布面积为1.20×106和1.332×106 km2 [10]。中国南方的喀斯特面积约为50万km2[11], 除去上述石漠化和非石漠化面积外,喀斯特山地森林系统(如广西的弄岗[12]、贵州的茂兰[13]、云南的西双版纳[14]和云南的石林[15])中,均发现很高比例的裸露岩体(图1)。喀斯特占世界陆地面积的12%15%[4],如果按照中国的石漠化及潜在石漠化的比例来推算,全球喀斯特生态系统中的含高露石比例的地域十分广阔。

2 露石上的生物殖居

露石上可殖居各种微生物。细菌(如硫细菌、硝化细菌和放线菌等)和藻类是岩石上最早出现的生物之一[16]。蓝藻(蓝细菌)中的黏球藻属、念珠藻属和真枝藻属,绿藻中小球藻属、软丝藻属和裂丝藻属等最为常见[17,18]。微生物可以石面生(Epilithic)、石内生(Endolithic)和石下生(Hypolithic)[19]。Matthes等[20]对加拿大温带地区一个地方的悬崖峭壁岩石表面生长层研究发现,蓝细菌、绿藻、地衣所占的比例分别为26%,3%和20%;李冰等[21]对一处喀斯特石漠藓类结皮调查后发现共有藓类植物7科9属13种,丛藓科(Pottiaceae)和牛舌藓科(Anomodontaceae)是优势科,平均生物量为1616 690 kg/hm2;田友萍等[22]对云南石林干燥的碳酸盐岩表面微生物研究发现,95%以上藻类为气生蓝藻,其胶被色泽鲜艳、宽厚而坚硬,是适宜高山岩石表面生活的典型进化特征。藻类和蓝藻多样性、丰富度与水分可利用性密切相关,真菌是需氧生物,一般占据岩石上层[23]。岩石微生物对有机质的分解促进了元素在石表面的生物地球化学循环[23]

藻类的孢子或细胞可以通过大气散播[24],且其表面形成的特殊淀粉体结构(Amyloid structures)[25]使藻类能有效黏附在石体表面,在一定量的水供应下进行光合作用和呼吸作用。气生藻类主要从大气中获取水,比如雨水、水汽、雾水,当水分不足时,藻类通过合成糖醇(Sugar alcohols)来调节细胞渗透压,维持细胞内稳态[26]。长期干旱时藻类会产生耐旱能力很强的孢子以应对干旱和延续生命[24];石生藻类则通过生物固氮、闪电固氮、大气沉降养分、溶蚀岩石矿物、有机质分解、有机氮的氨化等途径获取养分。温度、湿度、雨水、基质表面的物理特性和其他气候因子等因素共同影响岩石上藻类的分布[27]

地衣是真菌和藻类的共生体。蓝藻地衣和绿藻地衣分别利用大气中的水汽和液态水进行光合作用[24],且生长缓慢,一些叶状地衣10年间的生长量只有数厘米,被称为“时间色素”[28]。有研究表明,地衣的分布可能受到基质表面温度的限制,能耐受一定的极端高温,但也需要一段时间的低温来恢复生长和繁殖[27]。所以,在群落演替的初期,裸岩上最先迁入并定居的物种大多是藻类和地衣。

苔藓是继藻类、地衣演替阶段之后的石生群落,生境中温度、湿度、光照、基质持水性、岩石成分等的差异,可能导致不同的苔藓生活型出现[29]。在较平缓或微生境复杂的露石上,高等维管植物群落构成顶级群落,一般根系较浅的草本植物先出现,然后是乔灌木[30]

从群落演替的角度看,露石上的生物是一个演替早期植物群落。龚子同等[30]将岩石成土过程划分为3个阶段,每一阶段有其对应的殖居生物种类。藻类等着生的阶段属于第一阶段,也叫“岩漆”阶段,是积累营养元素的阶段;地衣着生阶段为第二阶段,是生物风化层形成发展的阶段;高等维管植物的殖居代表原始成土过程的终结。

徐海清等[31]对云南石林喀斯特山地原生植被—半湿润常绿阔叶林中附生维管植物的调查结果显示:林冠附生维管植物种类相对较少,只有8科9种附生维管束植物和5种苔藓植物;而林内岩石表面则有附生维管束植物16科17种,苔藓植物3种。

图2   露石与土壤斑块的组合形态示意

Fig.2   Illustration showing the combinations of rock outcrops and soi patches

朱习爱等[32]在云南石林的石漠化、人工林和自然林内的各60个2 m×2 m样方内分别记录到了15科21属21种、15科25属25种和28科39属41种维管束植物。进一步分析发现,维管束植物组成在3个系统间差异显著,表明不同的露石生境殖居差异巨大的植物群落。

3 露石对陆地生态系统的影响

如果将露石及其周围的土壤斑块看成“露石子系统”和“土壤子系统”,则可发现,露石影响最大的就是其邻近的“土壤子系统”。露石最直接的作用是承接降水(含穿透雨),大气干湿沉降[33,34],甚至是污染物[35],凝结雾水并携带沉降物中的N,P和K等输入土壤[36],Goransson等[37]把这种效应称之为斗效应(the funneling effect)。占据一定投影面积的石面上植居的生物能固定碳和氮[38],石面自身在不断溶蚀[39]为成土产物,接收并分解植根于土斑的各种植物的枯落物。到达石面的各类固体物质会以结皮形式附着于石面[40];或被石面上稀少的腐植土固定[41],但大量的集水将通过石面的各种裂隙和石—土界面进入地下管网系统;或通过石—土界面进入土壤,产生类似于干旱区的“斗效应”,影响土壤水分、养分和植物生长(生态水文效应),影响其生物群落的发展[38]。因此,“石体子系统”在一定程度上成为临近“土壤子系统”的潜在养分库,共同孕育出“石林”与“森林”共存的生态系统。但目前对调节裸露岩体的集水和径流输出的机理和生态水文效应尚不清楚。

按现有的一些数据推算,当裸露岩体占据一定的地面面积比例后,其石面汇集的水分、有机碳、营养物质的量是可观的。欧洲的森林氮沉降年通量变动于1404 200 kg/km2 [42];中国50个站点的年无机氮沉降总量变动于2604 820 kg/km2 [43];广东鼎湖山森林生态系统大气降水中的年总氮、钠、镁分别为384,21和12 kg/km2[44];云南滇池流域58月的总氮沉降(湿沉降)达190 kg/km2[45]。 随着工业活动的增强,大气沉降的数量还在增加[46]。目前还没有直接的测量结果来反映碳酸盐岩裸露岩体的石面生产能力,但如果参照火山(0.20.3 g/(m·a))[47]和陆地生物结皮(0.210.76 g/(m·a))[48]推算,数量也不会太低。

最近的研究发现,云南石林的石面径流输出约占降水收入的50%[49]。由此推理,在1∶ 1土石比的喀斯特生态系统中,其土面面积较无石系统减少50%,但余下的50%土面接收到的水分将相当于无石系统的150%。随后的检测还发现这些径流水中含有较高含量的有机碳和氮磷钾等内含物[50]。不同石面的径流输出变异很大,各种地貌类型内的裸露岩体千姿百态(图2),其物理特征(高度、坡度、粗糙度)、化学特征(可溶性)和生物学特征(附着物及其质量、腐殖土等)将是影响石面径流及其生态效果的重要因子。这些不同的“露石子系统”和“土壤子系统”形成丰富多彩的生态斑块组合,为植物的生长、生存提供了差异巨大的“生态位”空间,奠定了生物多样性基础。

地表千姿百态的露石在对邻近的土壤子系统及其生物产生影响的同时,还会对更大区域的水、土、物质转运产生重大影响,进而在更高级别的生态系统、甚至全球产生影响。现在已经有足够的证据显示,露石阻挡喀斯特系统地表径流,而将很高比例的水分导入地下管网,推动土壤失。应用137Cs证据计算得到的重庆槽谷区中地表流失的比例为74.55%,地下失比例为25.45%;而广西环江小流域土壤地面和地下流失的相对贡献率分别为12% 和88%[51],依据径流监测结果计算得到的广西峰丛洼地的山峰、垭口、山坡、山麓到峰丛洼地底部的土壤失占总流失量比例分别为92%,96%,79%,71%和39%[52]。这些巨大的数字差异或许与露石的出露比例有密切联系。土壤失后产生石漠化,已经成为中国南方十分严重的环境问题[9,10,53]。大气中的各种污染物也会通过这种富集作用,转入地下,污染地下水资源。同时,地下水的集中与溶蚀作用,还可能诱导地表塌陷等地质灾害[54]

4 未来研究

中国南方广泛分布的喀斯特露石生态系统,可以成为露石研究的模式生态系统。如果我们回头去重新审视广西的弄岗[12],贵州的茂兰[13],云南的西双版纳[14],云南的石林[15]等依然在森林掩盖下的喀斯特露石生态系统,会发现露石与植物,甚至高大的乔木“和谐相处”,维持着极高的植物多样性。我们不禁要问:露石在喀斯特生态系统中到底扮演了什么样的生态角色?我们或许忽略了太多的关于喀斯特露石—土壤—植物之间的相互作用规律。文献中能检索到喀斯特植物、土壤、土壤水分、生物地球化学[55]等的研究,也能从众多的石漠化治理实践者那直接或间接地领会到露石对植物生长的负面影响。少量研究关注到露石上生物的植居[38]、溶蚀[39],但无人关注其在露石生态系统中的潜在“正效应”。Li等[56]发现裸露岩体高度、方位均对土壤水分有影响。各喀斯特区的土壤分类中的黑色石灰土中有较高的有机质含量[30],这些研究结果均暗示了石面径流及其携带的内含物的生态水文效应,但缺乏足够的科学数据支撑,更不清楚其机制。

水是喀斯特过程中最重要的驱动因子[4,11],也是植物定居、更新与生长发育的关键因子[57]。稳定性同位素示踪结果显示,喀斯特植物能利用深层喀斯特水(与泉水具有相同同位素特征)和雾水[58],旱季尤为突出[59],但大部分的水分还是来源于土壤[60]。这些结果仍不能回答植物种群、群落与浅薄土壤水之间的关系,无法解释支撑南方喀斯特区域繁茂的森林植被及其丰富的植物物种多样性的水分格局。通过对距离露石不同远近、方位点位上的土壤水分含量变化,也推测到露石的汇水效应对土壤水分的影响[56],但缺乏足够定量的数据。未来需要扩展并完善露石径流水分研究,通过测定热带和亚热带露石汇集的径流水量,检测其内含物,系统地定量评价露石于热带、亚热带的汇水效应及其动态变化,及对土壤水分和营养物供应的影响,为阐明喀斯特生态系统演化机制做出。

植根于“土”的农耕文明遭遇到群体露石的石漠化问题时略显茫然,有人更将其看作类似于人类“癌症”的顽疾。继续沿着改造自然的大无畏指导思想努力去清除露石,砌石磊坎,植树遮掩(网上及各种政府报告中随处可见)或许能为我们带来新的植被,但付出是巨大的。当我们研究清楚露石—土壤—植物的关系后,或许我们的石漠化治理方略会展开新的视觉。

The authors have declared that no competing interests exist.


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DOI      URL      [本文引用: 1]      摘要

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