Distribution Characteristics of soil humus and fauna along the vertical natural belt of Lushan Mountain
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摘要: 沿庐山垂直自然带谱对常绿阔叶林(低海拔,L)、落叶常绿混交林(中海拔,M)和落叶阔叶林(高海拔,H)地面腐殖质及大中型土壤动物分布情况进行调查分析,使用调查数据探究了该垂直自然带土壤腐殖质与土壤动物分布特征。调查分析的腐殖质指标有剖面各发生层厚度、O-A过渡层厚度以及A层土壤团粒大小和有机质、总碳量、全磷、全氮、全碳含量和pH值等,土壤动物指标有类群数、个体密度和多样性指数。结果表明,L、M、H自然带地面腐殖质分别以细腐殖质(Mull)、半腐殖质(Moder)和粗腐殖质(Mor)为主;调查共获得隶属于3门8纲24目的大中型土壤动物2636头,其平均密度为2245.23 ind m−2,优势类群前气门亚目、甲螨亚目、弹尾目动物个体数分别占总个体数的38.87 %、25.36 %和13.37 %。皮尔逊相关性分析和典范对应分析(CCA)结果表明,土壤动物个体密度与A层土壤养分含量、类群数与O层厚度关系密切,而多样性指数既与A层土壤pH值、也与A层土壤养分含量及O-A过渡层厚度密切相关。土壤动物分布总体特点是优势类群在不同腐殖质组型土层广泛分布,而常见类群和稀有类群则分别与腐殖质组型中的OH层厚度、OL层厚度关系密切。Abstract: The distribution of surface humus and large and medium-sized soil fauna in evergreen broad-leaved forests (low altitude, L), deciduous evergreen mixed forests (medium altitude, M) and deciduous broad-leaved forests (high altitude, H) was investigated and analyzed along the altitudinal belt of Lushan Mountain, the distribution characteristics of soil humus and soil animals in the vertical natural zone were investigated by using survey data. The humus characters of investigation and analysis include the thickness of each occurrence layer, the thickness of O-A transition layers and aggregates size of A-horizon and organic matter, total carbon content, total phosphorus, total nitrogen, total carbon content and pH value of A layer. Soil fauna characters included the number of biological groups, individual density and diversity index. The results showed that the surface humus of L, M and H natural zones were mainly fine humus (Mull), semi-humus (Moder) and coarse humus (Mor), respectively. A total of 2636 large and medium-sized soil fauna were identified, belonging to 24 orders in 8 classes under 3 phyla, in which the average density was 2245.23 ind m−2. Among them, the dominant group was Prostigmata, Oribatida, Collembola, and the number of individuals in the dominant groups was 38.87%, 25.36% and 13.37% respectively. The pearson Correlation and CCA analysis showed that the individual density of soil fauna was closely related to soil nutrient content in A-horizon and the number of biological group was highly associated with the thickness of O horizon. The diversity index was closely connected not only to soil pH of A-horizon but also to soil nutrient content in A-horizon and the thickness of O-A transition layer. The overall characteristics of soil fauna distribution are that dominant groups are widely distributed in different humus groups, while the common groups were in close relation with the thickness of OH layer, but the rare groups had an intimate connection with the thickness of OL layer.
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Key words:
- Community structure /
- Humus form /
- Altitude /
- Fauna /
- Diversity
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图 2 土壤动物与腐殖质组型典范对应分析排序图
1:前气门亚目 Prostigmata;2:甲螨亚目Oribatida;3:弹尾目Collembola;4:双翅目幼虫Diptera larvae;5:膜翅目Hymenoptera;6:中气门亚目Mesostigmata;7:鼠妇科Porcellio;8:蜘蛛目Araneae;9:鞘翅目幼虫Coleoptera larva;10:倍足纲Diplopoda;11:巨蚓科Megascolecidae;12:鳞翅目幼虫Lepidoptera larva
Figure 2. Canonical correspondence analysis ordination of soil fauna and humus form
表 1 庐山垂直自然带及样地的基本情况
Table 1. The basic situation of the vertical natural belt and sites of Lushan
样地
Site经纬度
Longitude and
latitude海拔
Altitude
(m)当日气温
Temperature
(℃)年降水量
Annual Precipitation
(mm)坡向
Aspect of slope岩性
Lithology土壤类型
Soil type植被类型
Vegetation主要植物组成
Main plantcomposition大月山(H) 29°33′42″N115°00′E 1320 18 1933 阴坡 片麻岩 山地棕壤 落叶阔叶林 山檀(Santalum paniculatum) 黄龙寺(M) 29°33′6″N115°57′45″E 910 20 2208 阴坡 含砾石英砂岩 山地黄棕壤 落叶常绿混交林 阴香(Cinnamomum burmanni(Nees et T.
Nees)blume),白簕(Acanthopanax
trifoliatus(L.)Merr)白鹿洞(L) 29°21′35″N116°2′21″E 120 25 1124 阳坡 花岗岩 红壤 常绿林 青冈(Quercus glauca Thunb) 表 2 不同海拔下腐殖质组型的指标(平均值 ± 标准误)
Table 2. Indices of humus form at different altitudes(Mean ± SE)
指标
Index1320 m 910 m 120 m OL层厚度(mm) 18.33 ± 1.67 c 26.67 ± 1.67 b 50.00 ± 5.00 a OF层厚度(mm) 16.00 ± 1.00 a 17.67 ± 1.45 a 9.00 ± 1.00 b OH层厚度(mm) 18.33 ± 1.67 a 13.33 ± 1.67 a 0.00 ± 0.00 b O-A过渡层厚度(mm) 2.53 ± 0.15 b 6.50 ± 0.29 a 1.77 ± 0.15 c A层厚度(mm) 58.33 ± 4.23 a 55.00 ± 5.77 ab 37.33 ± 1.20 b A层团粒大小(mm) 0.77 ± 0.09 c 3.13 ± 0.41 b 6.67 ± 0.88 a A层pH值 3.90 ± 0.07 a 4.52 ± 0.14 c 5.02 ± 0.09 b A层 TN(g kg−1) 7.91 ± 1.00 b 5.74 ± 0.17 a 2.34 ± 0.46 a A层 TC(g kg−1) 112.60 ± 16.85 b 66.94 ± 1.76 a 28.95 ± 7.31 a A层 TP(g kg−1) 1.09 ± 0.12 b 0.67 ± 0.03 a 0.46 ± 0.07 b A层 OM(g kg−1) 225.33 ± 40.29 b 128.58 ± 17.24 a 50.94 ± 17.53 a A层 Hu(g kg−1) 123.31 ± 21.48 a 81.56 ± 5.20 a 28.57 ± 9.91 b A层 C/N 14.01 ± 0.41 a 11.69 ± 0.32 b 11.74 ± 0.82 b 注:OL层:完整未分解凋落物;OF层:半分解破碎凋落物;OH层:完全分解的凋落物及动物粪便;A层:有机-矿物层;TN全氮:Total nitrogen;TC:总碳 Total carbon;TP:总磷 Total phosphorus;OM:有机质Organic matter;Hu:腐殖质总碳量;C/N:碳氮比Carbon-nitrogen ratio。表中同行带相同字母者差异不显著,不同字母者差异显著(P < 0.05, LSD)。同下。 表 3 土壤动物群落组成(平均值 ± 标准误)
Table 3. Soil fauna community composition(Mean ± SE)
类群
Group1320 m 910 m 120 m 总和
Sum丰度
Percent密度(ind m−2)
Density密度(ind m−2)
Density密度(ind m−2)
Density密度(ind m−2)
Density% 前气门亚目 Prostigmata 353.3 ± 26.9 1939.8 ± 10.3 325.0 ± 210.4 2618.1 ± 247.6 38.87 甲螨亚目 Oribatida 597.2 ± 47.4 482.2 ± 34.0 628.8 ± 279.5 1708.2 ± 360.9 25.36 弹尾目 Collembola 142.2 ± 41.4 277.8 ± 23.5 480.5 ± 253.3 900.5 ± 318.2 13.37 中气门亚目 Mesostigmata 122.2 ± 122.2 150.0 ± 150.0 27.8 ± 9.6 300.0 ± 281.8 4.45 双翅目幼虫 Diptera larva 133.3 ± 31.3 135.5 ± 27.5 26.7 ± 5.7 295.5 ± 64.5 4.39 鼠妇科 Porcellio 100.0 ± 63.2 13.9 ± 2.8 14.8 ± 3.7 128.7 ± 69.5 1.91 膜翅目 Hymenoptera 14.8 ± 3.7 28.9 ± 8.2 80.0 ± 46.7 123.7 ± 58.6 1.84 蜘蛛目 Araneae 27.8 ± 13.2 20.4 ± 3.4 33.3 ± 4.5 81.5 ± 21.1 1.21 鞘翅目幼虫 Coleoptera larva 37.8 ± 14.3 18.5 ± 7.4 56.3 ± 21.7 0.84 巨蚓科 Megascolecidae 33.3 ± 33.3 22.2 ± 11.1 55.5 ± 55.5 0.82 白蚁科 Termitidae 44.4 ± 44.4 11.1 ± 11.1 55.5 ± 55.5 0.82 石蜈蚣目 Lithobiomorpha 24.4 ± 5.4 11.1 ± 11.1 11.1 ± 11.1 46.6 ± 27.6 0.69 地蜈蚣目 Geophilomorpha 33.3 ± 11.1 11.1 ± 11.1 44.4 ± 22.2 0.66 综合目 Symphyla 11.1 ± 11.1 11.1 ± 11.1 22.2 ± 6.4 44.4 ± 28.6 0.66 直翅目 Orthoptera 11.1 ± 11.1 33.3 ± 33.3 44.4 ± 44.4 0.66 隐翅虫科 Staphylinidae 17.8 ± 6.7 11.1 ± 11.1 11.1 ± 11.1 40.0 ± 28.9 0.59 倍足纲 Diplopoda 25.9 ± 14.8 11.1 ± 11.1 37.0 ± 25.9 0.55 拟蝎目 Pseudoscorpiones 11.1 ± 11.1 22.2 ± 22.2 33.3 ± 33.3 0.49 大蜈蚣目 Scolopendrida 11.1 ± 11.1 11.1 ± 11.1 22.2 ± 22.2 0.33 半翅目 Hemiptera 11.1 ± 11.1 11.1 ± 11.1 22.2 ± 22.2 0.33 鳞翅目幼虫 Lepidoptera larva 11.1 ± 11.1 11.1 ± 11.1 0.16 端足目 Amphipoda 11.1 ± 11.1 11.1 ± 11.1 0.16 蜚蠊目 Blattodea 11.1 ± 11.1 11.1 ± 11.1 0.16 革翅目 Dermaptera 11.1 ± 11.1 11.1 ± 11.1 0.16 柄眼目 Stylommatophora 11.1 ± 11.1 11.1 ± 11.1 0.16 腹足目 Gastropoda 11.1 ± 11.1 11.1 ± 11.1 0.16 鞘翅目成虫 Coleoptera imago 11.1 ± 11.1 11.1 ± 11.1 0.16 总密度 Total density 1696.6 ± 261.0 3300.1 ± 461.6 1739.0 ± 620.6 6735.7 ± 940.2 100 类群数 Group nullmber 19 23 16 27 注:优势类群占总个体数10%以上,常见类群占总个体数1% ~ 10%,稀有类群占总个体数1%以下。 表 4 不同海拔下土壤动物多样性指标(平均值 ± 标准误)
Table 4. Indices of soil fauna diversity at different altitudes (Mena ± SE)
多样性指数
Diversity index1320 m 910 m 120 m S 9.67 ± 0.92 a 9.67 ± 0.61 a 8.00 ± 1.00 a H′ 1.62 ± 0.04 a 1.26 ± 0.10 b 1.50 ± 0.09 a J 0.73 ± 0.03 ab 0.57 ± 0.06 b 0.75 ± 0.07 a C 0.26 ± 0.02 b 0.42 ± 0.04 a 0.29 ± 0.04 b 注:S:类群数;H′:多样性指数;J:均匀度指数;C:优势度指数。同下。 表 5 土壤动物与腐殖质组型指标之间的Pearson相关系数
Table 5. Correlation coefficients between soil fauna and humus morphological index
指标
Index土壤动物个体密度
Number of density土壤动物类群数
Number of groupH′ J C A层pH值 −0.432 −0.084 0.517* 0.359 −0.492* A层 TN 0.439 0.130 −0.576* −0.448 0.541* A层 TC 0.347 0.132 −0.495* −0.396 0.470* A层 TP 0.539* 0.303 −0.612** −0.590** 0.594** A层 Hu 0.202 0.348 0.020 −0.244 0.032 A层 OM 0.276 0.010 −0.519* −0.320 0.441 A层 C/N 0.118 0.384 0.410 −0.015 −0.338 A层 N/P 0.507* 0.329 −0.369 −0.478* 0.421 A层 C/P 0.458 0.422 −0.121 −0.394 0.196 OL层厚度 −0.001 −0.796* 0.125 0.526 −0.300 OF层厚度 0.141 0.646 −0.406 −0.626 0.517 OH层厚度 0.110 0.867** −0.103 −0.515 0.245 O-A过渡层厚度 0.488 0.331 −0.865** −0.814** 0.916** A层厚度 0.114 0.584 −0.086 −0.373 0.186 A层团粒大小 0.054 −0.658 −0.088 0.256 −0.045 注:**在0.01 水平(双尾)上显著相关; * 在 0.05 水平(双尾)上显著相关 -
[1] Zanella A, Jabiol B, Ponge J F, et al. A European morpho-functional classification of humus forms[J]. Geoderma, 2011, 164(3): 138 − 145. [2] 全国自然科学名词审定委员会. 林学名词[M]. 科学出版社, 1989. [3] Ponge J F. Humus forms in terrestrial ecosystems: a framework to biodiversity[J]. Soil Biology and Biochemistry, 2003, 35(7): 935 − 945. [4] Zanella A, Berg B, Ponge J F, et al. Humusica 1, article 2: Essential bases-functional considerations[J]. Applied Soil Ecology, 2018, 122: 22 − 41. doi: 10.1016/j.apsoil.2017.07.010 [5] Zanella A, Ponge J F, Brionesc J I M. Humusica 1, article 8: Terrestrial humus systems and forms – biological activity and soil aggregates, space-time dynamics[J]. Applied Soil Ecology, 2018, 122: 103 − 137. doi: 10.1016/j.apsoil.2017.07.020 [6] 路有成, 王宗英. 九华山土壤动物的垂直分布[J]. 地理研究, 1994, 13(2): 74 − 81. [7] Nicolas B, Francois G. Structural relationships among vegetation, soil fauna and humus form in a subalpine forest ecosystem: a hierarchical multiple factor analysis (HMFA)[J]. Pedobiologia, 2012, 55(6): 321 − 334. doi: 10.1016/j.pedobi.2012.06.004 [8] Sandrine S, Nadia A, Lorenzo F, et al. Relationships between soil fauna communities and humus forms: Response to forest dynamics and solar radiation[J]. Soil Biology and Biochemistry, 2008, 40(7): 1707 − 1715. doi: 10.1016/j.soilbio.2008.02.007 [9] Waez-mousavi S M. Humus systems in the Caspian Hyrcanian temperate forests[J]. Applied Soil Ecology, 2018, 123: 664 − 667. doi: 10.1016/j.apsoil.2017.09.022 [10] Michaël A, Jean T, Matthieu C, et al. Forest humus forms as a playground for studying aboveground-belowground relationships: Part 2, a case study along the dynamics of a broadleaved plain forest ecosystem[J]. Applied Soil Ecology, 2018, 123: 398 − 408. doi: 10.1016/j.apsoil.2017.09.005 [11] Labaz B, Galka B, Bogacz A, et al. Factors influencing humus forms and forest litter properties in the mid-mountains under temperate climate of southwestern Poland[J]. Geoderma, 2014, 230-231: 265 − 273. [12] Bayrandvand M, Kooch Y, Hosseini S M, et al. Humus forms in relation to altiutude and forest type in the northern mountainous regions of Iran[J]. Forest Ecology and Management, 2017, 385: 78 − 86. doi: 10.1016/j.foreco.2016.11.035 [13] 周育臻, 吴鹏飞. 贡嘎山东坡森林小型土壤节肢动物群落多样性与时空分布[J]. 生态学杂志, 2020, 39(2): 586 − 599. [14] 高宝嘉. 雾灵山森林植物与节肢动物群落结构及多样性研究[D]. 北京: 北京林业大学, 2005. [15] 徐帅博, 李艳红, 朱连奇, 等. 山地土壤动物群落垂直分布格局研究进展[J]. 河南大学学报(自然科学版), 2020, 50(1): 19 − 28. [16] Kathman R D, Cross S F. Ecological distribution of moss-dwelling tardigrade on Vancouver Island. British Columbia, Canada[J]. Canadian Journal of Zoology, 1991, 69: 122 − 129. doi: 10.1139/z91-018 [17] Ohsawa M. Latitudinal comparison of altitudinal changes in forest structure, leaf-type, and species richness in humid monsoon Asia[J]. Vegetatio, 1995, 121: 3 − 10. doi: 10.1007/BF00044667 [18] 王振中, 李忠武, 张友梅. 庐山人工针叶林土壤动物群落调查[J]. 湖南师范大学自然科学学报, 1998, 12(4): 83 − 88. [19] 陈小鸟, 由文辉, 易 兰. 浙江天童太白山不同海拔土壤动物的群落结构[J]. 生态学杂志, 2009, 28(2): 270 − 276. [20] Grytnes J A. Species-richness patterns of vascular plants along seven altitudinal transects in Norway[J]. Ecography, 2003, 26(3): 291 − 300. [21] 张金泉. 庐山植被的垂直分带[J]. 华南师院学报(自然科学版), 1982, (1): 1 − 14. [22] 汪权方, 王宗英. 庐山土壤动物群落结构研究[J]. 安徽师范大学学报(自然科学版), 2003, 26(2): 180 − 185. [23] 廖丽琴, 刘苑秋, 孔凡前, 等. 庐山冬季土壤动物群落及功能群对毛竹扩张的响应[J]. 江西农业大学学报, 2017, 39(4): 721 − 730. [24] 刘信中, 王 琅. 江西省庐山自然保护区生物多样性考察与研究[M]. 北京, 科学出版社, 2010. [25] 青木淳一. 土壤动物[M]. 东京: 北隆馆, 1973. [26] 尹文英. 中国土壤动物检索图鉴[M]. 北京: 科学出版社, 1998. [27] Zanella A, Ponge J F, Jabiol B, et al. Humusica 1, article 5: Terrestrial humus systems and forms-Keys of classification of humus systems and forms[J]. Applied Soil Ecology, 2018, 122: 75 − 86. [28] Pielou E C. Ecology Diversity[M]. New York: JohnWieley & Sons, 1975. [29] 尹文英. 中国土壤动物[M]. 北京: 科学出版社, 2000. [30] Mohammad B, Yahya K, Giorgio A. Classification of humus forms in Caspian Hyrcanian mixed forests ecoregion (Iran): Comparison between two classification methods[J]. Catena, 2018, 165: 390 − 397. doi: 10.1016/j.catena.2018.02.021 [31] Beate S, Søren M, Kristiansen B, et al. Dynamic oak-scrub to forest succession: Effects of management on understorey vegetation, humus forms and soils[J]. Forest Ecology and Management, 2005, 211(3): 318 − 328. [32] Berg B, Meentemeyer V. Litter fall in some European coniferous forests as dependent on climate: A synthesis[J]. Canadian Journal of Forest Research, 2001, 31: 292 − 301. doi: 10.1139/x00-172 [33] Akselsson C, Berg B, Meentemeyer V, et al. Carbon sequestration rates in organic layers of boreal and temperate forest soils-Sweden as a case study[J]. Global Ecology and Biogeography, 2005, 14: 77 − 84. [34] Berg B, Ekbohm G, Johansson M B, et al. Maximum decomposition limits of forest litter types: a synthesis[J]. Canadian journal of Botany, 1996, 74: 659 − 672. doi: 10.1139/b96-084 [35] Berg B, Johanssonm B, Nilssonå, et al. Sequestration of carbon in the humus layer of Swedish forests Å direct measurements[J]. Canadian Journal of Forest Research, 2009, 39: 962 − 975. doi: 10.1139/X09-022 [36] Lieberman D, Lieberman M, Peralta R, et al. Tropical forest structure and compositionon a large-scale altitudinal gradientin in Costa Rica[J]. Journal of Ecology, 1996, 84(2): 137 − 152. [37] Xu G R, Zhang Y X, Zhang S, et al. Biodiversity associations of soil fauna and plants depend on plant life form and are accounted for by rare taxa along an elevational gradient[J]. Soil Biology and Biochemistry, 2020, 140: 107640. doi: 10.1016/j.soilbio.2019.107640 [38] 肖能文, 刘向辉, 戈 峰, 等. 高黎贡山自然保护区大型土壤动物群落特征[J]. 生态学报, 2009, 29(7): 3576 − 3584. doi: 10.3321/j.issn:1000-0933.2009.07.016 [39] Whittaker R H, Niering W A. Vegetation of the Santa Catalina Mountains, Arizona.Ⅴ.Biomass, production, and diversity along the elevation gradient[J]. Ecology, 1975, 56: 771 − 790. doi: 10.2307/1936291 [40] 王邵军, 阮宏华, 汪家社, 等. 武夷山典型植被类型土壤动物群落的结构特征[J]. 生态学报, 2010, 30(19): 5174 − 5184. [41] 陈颖彪, 殷秀琴. 凉水地区不同林型土壤动物群落研究[J]. 上海师范大学学报(自然科学版), 2000, 29(2): 79 − 84. [42] Frak E, Ponge J F. The influence of altitude on the distribution of subterranean organs and humus components in Vaccinium myrtillus carpets[J]. Journal of Vegetation Science, 2002, 13: 17 − 26. doi: 10.1658/1100-9233(2002)013[0017:TIOAOT]2.0.CO;2 [43] Sophya S. Collembolan communities of the Ubsu -Nur Basin and adjacent mountains (Russia, Tuva)[J]. Pedobiologia, 2003, 47(4): 341 − 356. [44] 辛未冬, 殷秀琴, 秦丽杰, 等. 小兴安岭森林生态系统中小型土壤动物生态地理分布特征[J]. 地理科学, 2009, 29(1): 129 − 133. doi: 10.3969/j.issn.1000-0690.2009.01.021 [45] 岳 明, 张林静, 党高弟, 等. 佛坪自然保护区植物群落物种多样性与海拔梯度的关系[J]. 地理科学, 2002, 22(3): 349 − 354. doi: 10.3969/j.issn.1000-0690.2002.03.016 [46] 严 莹, 李 恺, 方 燕. 浙江百山祖自然保护区不同海拔土壤动物群落结构及季节动态[J]. 生态学杂志, 2010, 29(9): ©1754 − 1767. [47] Gabin C, Apolline A, Françoise W. Techno-moder: A proposal for a new morpho-functional humus form developing on Technosols revealed by micromorphology[J]. Geoderma, 2020, 375: 114526. doi: 10.1016/j.geoderma.2020.114526 [48] Salmon S. Changes in humus forms, soil invertebrate communities and soil functioning with forest dynamics[J]. Applied Soil Ecology, 2018, 123(2): 345 − 354.