中高山岩溶区典型群落土壤–植物–凋落物连续体碳氮磷养分特征及影响因素

史佳良; 陶兰初; 罗平; 贺思腾; 湛方栋

doi:10.11932/karst20250606

中高山岩溶区典型群落土壤–植物–凋落物连续体碳氮磷养分特征及影响因素

doi: 10.11932/karst20250606

史佳良^{1, 3, 4,},
陶兰初^{1, 3, 4, ,},
罗平²,
贺思腾²,
湛方栋²

1.
中国地质调查局昆明自然资源综合调查中心, 云南昆明 650111
2.
云南农业大学资源与环境学院, 云南昆明 650201
3.
中国地质学会西南山地生态地质演化与保护修复创新基地, 云南昆明 650111
4.
自然资源部自然生态系统碳汇工程技术创新中心, 云南昆明 650111

基金项目: 中国地质调查局项目（DD20230483，DD20230098)

详细信息

作者简介:
史佳良（1990－），男，工程师，硕士研究生，主要从事生态地质与石漠化研究。E-mail：376852612@qq.com

通讯作者:
陶兰初（1989－），男，工程师，硕士研究生，主要从事环境地球化学研究。E-mail：690286149@qq.com。

中图分类号: S714
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出版历程
- 收稿日期: 2024-10-29
- 录用日期: 2025-08-27
- 修回日期: 2025-07-15
- 刊出日期: 2025-12-25

Characteristics and influencing factors of carbon, nitrogen, and phosphorus nutrients in the soil-plant-litter continuum of typical communities in mid-high mountain karst areas

SHI Jialiang^{1, 3, 4
,},
TAO Lanchu^{1, 3, 4
, ,},
LUO Ping²,
HE Siteng²,
ZHAN Fangdong²

1.
Kunming General Survey of Natural Resources Center, China Geological Survey, Kunming, Yunnan 650111, China
2.
College of Resources and Environment, Yunnan Agricultural University, Kunming, Yunnan 650201, China
3.
Innovation Base for Ecogeological Evolution, Protection and Restoration of Southwest Mountainous Areas, Geological Society of China, Kunming, Yunnan 650111, China
4.
Technology Innovation Center for Natural Ecosystem Carbon Sink, Ministry of Natural Resources, Kunming, Yunnan 650111, China

摘要

摘要: 中高山岩溶区具有独特的地理和环境特征，生态系统脆弱，研究土壤–植物–凋落物的物质循环和能量流动在各个典型植被群落之间的变化，对于理解该地区的生态恢复和可持续发展具有重要意义。文章以滇西北中高山岩溶石漠化区的5种典型群落类型（草地、灌草、乔灌草–柏、纯松、乔灌草–松）选取的23个不同群落共69块样地为研究对象，分析土壤、植物和凋落物的碳（C）、氮（N）、磷（P）含量与化学计量特征，以及环境因素的影响。结果表明：（1）不同群落类型土壤C、N、P含量均值为7.46~43.25 g·kg⁻¹、1.05~2.96 g·kg⁻¹、0.39~0.53g·kg⁻¹，土壤C/N、C/P、N/P计量比为8.82~23.25、21.74~105.25、2.89~6.33；植物C、N、P含量为373.79~622.87 g·kg⁻¹、10.47~15.56 g·kg⁻¹、1.43~2.14 g·kg⁻¹，植物C/N、C/P、N/P计量比为27.10~65.13、272.85~624.26、7.23~10.72；凋落物C、N、P含量为334.93~491.73 g·kg⁻¹、7.00~16.61 g·kg⁻¹、1.16~1.53 g·kg⁻¹，凋落物C/N、C/P、N/P计量比为23.70~72.17、238.49~499.18、7.61~17.49，不同群落类型土壤、植物和凋落物的C、N、P含量及化学计量比差异显著；（2）土壤与植物的C、N养分基本上表现出随群落类型由草地到乔木逐渐丰富的趋势，乔灌草–松、纯松、灌草阶段普遍大于草地和乔灌草–柏阶段，而凋落物则与之相反；土壤普遍缺P，但植物与凋落物中未表现出缺P；（3）土壤C/N值、C/P值、N/P值在纯松、乔灌草–松、灌草类型显著大于乔灌草–柏和草地；植物与凋落物的C/N值、C/P值乔灌草–柏类型显著大于其他类型；（4）土壤、植物和凋落物之间的养分含量呈现相关关系，群落类型、海拔、石漠化程度等环境因子对化学计量特征有显著影响，且各因子间相关性复杂，影响重要性排序为海拔＞生态修复措施＞石漠化程度＞群落类型＞土壤厚度＞植被覆盖度；（5）中高山岩溶区植被生长受到N、P的共同制约，乔木群落相较草本与矮灌群落，更多受到N素的制约。
- 岩溶区 /
- 典型群落类型 /
- 土壤–植物–凋落物连续体 /
- 碳氮磷化学计量特征 /
- 影响因子
Abstract: This study focuses on the mid-high mountain karst rocky desertification areas in northwestern Yunnan, aiming to elucidate the stoichiometric characteristics and influencing mechanisms of carbon (C), nitrogen (N), and phosphorus (P) within the soil-plant-litter continuum in this region. It is of great significance for an in-depth understanding of ecosystem cycles in mid-high mountain karst areas and provides theoretical support for ecological restoration efforts.The study area is located in Shuhe town and Jiuhe township, south of Yulong Snow Mountain, on the southeastern edge of the Qinghai-Xizang Plateau. It lies within the transition zone between China's first and second terrain steps. The region features complex terrain with numerous high mountains and valleys, and an altitude ranging from 2,544 m to 3,090 m. Carbonate rocks are widely distributed, with limestone as the main soil-forming parent material, resulting in an extremely slow rate of soil formation through weathering; The predominant soil type is brown soil. The area experiences a plateau monsoon climate characterized by distinct dry and wet seasons, an annual average temperature of 13℃ to 20 ℃, and an annual average rainfall of 950 mm to1200 mm, over 85% of which occurs between May and October. Solar radiation remains strong throughout the year. Vegetation is characterized by xerophytism, lithophytism, and calciphilia, with key species including Arundinella hookeri Munro ex Keng, Tripogon yunnanensis J. L. Yang ex S. M. Phillips & S. L., Festuca ovina L., Lespedeza forrestii Schindl., Rhododendron racemosum Franch., Quercus guyavifolia H. Lév., Cotoneaster microphyllus Wall. ex Lindl., Pinus yunnanensis Franch., Juniperus formosana Hayata, Cupressus duclouxiana Hickel.Targeting the ecological vulnerability of mid-high mountain karst areas and addressing existing research gaps, this study analyzes the C, N, and P contents and stoichiometric characteristics of soil, plants, and litter under different vegetation community types. It further explores the impacts of community types and environmental factors on nutrient cycling. This study focuses on answering two core questions: first, the variation patterns of C, N, and P contents and ecological stoichiometric characteristics in the soil-plant-litter continuum; Second, the specific effects of vegetation community types on C, N, and P contents within this continuum.From July to August 2023, the survey was conducted in the study area encompassing five typical community types: grassland, shrub-grass, arbor-shrub-grass-cypress, pure pine, and arbor-shrub-grass-pine. The grassland type includes three communities, while the shrub-grass, arbor-shrub-grass-cypress, pure pine, and arbor-shrub-grass-pine types each included five communities, totaling 23 communities. For each community, three parallel sample plots were selected, resulting in a total of 69 sample plots. In each sample plot, a standard 10 m×10 m quadrat was established, along with a representative 5 m×5 m shrub quadrat and five 1 m×1 m herb quadrats arranged using the plum-point method. Indicators such as geographical location, altitude, community type, ecological restoration measures, soil thickness, vegetation coverage, rock exposure rate, and degree of rocky desertification were recorded. Community diversity was assessed using indices such as species richness and the Shannon-Wiener diversity index. Samples were analyzed for C, N, and P contents and stoichiometric characteristics of soil, plants, and litter, as well as the influence of environmental factors.The research results show that the average contents of soil C, N, and P across different community types were 7.46 to 43.25 g·kg⁻¹, 1.05 to 2.96 g·kg⁻¹, and 0.39 to 0.53 g·kg⁻¹, respectively. The soil C/N, C/P, and N/P ratios ranged from 8.82 to 23.25, 21.74 to 105.25, and 2.89 to 6.33, respectively. Plant C, N, and P contents were 373.79 to 622.87 g·kg⁻¹, 10.47 to 15.56 g·kg⁻¹, and 1.43 to 2.14 g·kg⁻¹, with plant C/N, C/P, and N/P ratios of 27.10 to 65.13, 272.85 to 624.26, and 7.23 to 10.72, respectively. Litter C, N, and P contents were 334.93 to 491.73 g·kg⁻¹, 7.00 to 16.61 g·kg⁻¹, and 1.16 to1.53 g·kg⁻¹, with litter C/N, C/P, and N/P ratios of 23.70 to 72.17, 238.49 to 499.18, and 7.61 to 17.49, respectively. Significant differences were observed in the C, N, P contents and stoichiometric ratios of soil, plants, and litter among different community types. Soil and plant C and N nutrients generally showed a trend of gradual enrichment as community types transitioned from grassland to arbor, Specifically, the arbor-shrub-grass-pine, pure pine, and shrub-grass stages generally showed higher nutrient levels than the grassland and arbor-shrub-grass-cypress stages, while litter exhibited the opposite trend. Soil was generally deficient in P, but plants and litter did not exhibit signs of P deficiency. In terms of stoichiometric ratios, and soil C/N, C/P, and N/P ratios in pure pine, arbor-shrub-grass-pine, and shrub-grass types were significantly higher than those in arbor-shrub-grass-cypress and grassland types. Conversely, plant and litter C/N and C/P ratios in arbor-shrub-grass-cypress type were significantly higher than those in other types. There were widespread correlations in nutrient contents among soil, plants, and litter. Environmental factors such as community type, altitude, and degree of rocky desertification had significant impacts on stoichiometric characteristics, with complex correlations among factors. The relative importance of these influences was ranked as follows, altitude > ecological restoration measures > degree of rocky desertification > community type > soil thickness > vegetation coverage.Vegetation growth in mid-high mountain karst areas is jointly limited by N and P, with the degree of limitation varying among different community types. Compared with herbaceous and low shrub communities, arbor communities are more strongly constrained by N.
- karst area /
- typical community types /
- soil-plant-litter continuum /
- stoichiometric characteristics of carbon, nitrogen and phosphorus /
- influencing factors

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图 1 典型群落类型土壤-植物-凋落物C、N、P含量

注：图中S-C为土壤C含量，S-N为土壤N含量，S-P为土壤P含量；P-C为植物C含量，P-N为植物N含量，P-P为植物P含量；L-C为凋落物C含量，L-N为凋落物N含量，L-P为凋落物P含量。

Figure 1. Contents of C, N, and P of soil-plant-litter at different typical community types

Note: S-C represents soil carbon content, S-N represents soil nitrogen content, and S-P represents soil phosphorus content. P-C represents plant carbon content, P-N represents plant nitrogen content, and P-P represents plant phosphorus content. L-C represents litter carbon content, L-N represents litter nitrogen content, and L-P represents litter phosphorus content.

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图 2 典型群落类型土壤–植物–凋落物C/N、C/P、N/P比值

注：图中S-C/N为土壤碳氮比，S-C/P为土壤碳磷比，S-N/P为土壤氮磷比；P-C/N为植物碳氮比，P-C/P为植物碳磷比，P-N/P为植物氮磷比；L-C/N为凋落物碳氮比，L-C/P为凋落物碳磷比，L-N/P为凋落物氮磷比。

Figure 2. Ratios of C/N, C/P, and N/P of soil-plant-litter at different typical community types

Note: S-C/N represents the soil carbon-nitrogen ratio, S-C/P represents the soil carbon-phosphorus ratio, and S-N/P represents the soil nitrogen-phosphorus ratio. P-C/N represents the plant carbon-nitrogen ratio, P-C/P represents the plant carbon-phosphorus ratio, and P-N/P represents the plant nitrogen-phosphorus ratio. L-C/N represents the litter carbon-nitrogen ratio, L-C/P represents the litter carbon-phosphorus ratio, and L-N/P represents the litter nitrogen-phosphorus ratio.

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图 3 土壤–植物–凋落物C、N、P化学计量特征与环境因子的相关性

注：图中CMT为群落类型，ERM为生态修复措施，AL为海拔高度，DoRD为石漠化程度，RER为岩石裸露率，VC为植被覆盖度，ST为土壤厚度，R为丰富度指数，H为香农–威尔指数，E为均匀度指数，D为优势度指数；“*”表示显著性P＜0.05，“**”表示显著性P＜0.01，“***”表示显著性P＜0.001，“****”表示显著性P＜0.0001。

Figure 3. Correlation between the stoichiometric characteristics of C, N, and P in soil-plant-litter and environmental factors

Note: CMT represents community type, ERM represents ecological restoration measure, AL represents altitude, DoRD represents degree of rocky desertification, RER represents rock exposure rate, VC represents vegetation coverage, ST represents soil thickness, R represents richness index, H represents Shannon-Wiener index, E represents evenness index, and D represents dominance index. “*” indicates significance with P < 0.05, “**” indicates significance with P < 0.01, “***” indicates significance with P < 0.001, and “****” indicates significance with P < 0.0001.

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图 4 土壤-植物-凋落物C、N、P及化学计量比与主要环境因子的RDA分析

注：图中CMT为群落类型，ERM为生态修复措施，AL为海拔高度，DoRD为石漠化程度，RER为岩石裸露率，VC为植被覆盖度，ST为土壤厚度。

Figure 4. RDA analysis of soil-plant-litter C, N, P and stoichiometric ratios with main environmental factors

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表 1 样点概况信息

Table 1. Basic information of sample plots

群落名称	群落类型	经度	纬度	海拔/m	石漠化程度	生态修复措施	pH	岩石裸露率/%	植被覆盖度/%	土壤厚度/cm
小叶栒子–旱茅群落	草地1	100°10′32.36″E	26°39′56.56″N	2612	中度	无措施	6.34±0.02ab	60	28	6
矮生胡枝子–西南野古草群落	草地2	100°10′31.68″E	26°39′55.79″N	2677	中度	无措施	6.18±0.10b	42	31	8
西南野古草群落	草地3	100°10′17.72″E	26°39′24.54″N	2648	重度	无措施	6.08±0.03b	52	25	15
刺叶高山栎–西南野古草群落	灌草1	100°16′59.78″E	27°02′15.11″N	3049	轻度	无措施	6.61±0.06a	35	51	15
刺叶高山栎–狼毒群落	灌草2	100°16′59.82″E	27°02′13.18″N	3040	中度	无措施	6.02±0.13b	32	40	8
帽斗栎–西南野古草群落	灌草3	100°17′22.74″E	27°01′42.11″N	3090	重度	无措施	6.33±0.17ab	58	40	10
帽斗栎–西南野古草群落	灌草4	100°18′48.30″E	27°00′52.49″N	2795	重度	无措施	6.29±0.11ab	68	38	8
帽斗栎–灯心草群落	灌草5	100°18′50.58″E	27°00′49.19″N	2794	重度	无措施	6.45±0.08ab	65	35	8
刺柏–腋花杜鹃–西南野古草群落	乔灌草–柏1	100°10′13.08″E	26°39′36.89″N	2669	重度	植树造林	5.80±0.23c	51	29	15
刺柏–小叶栒子–西南野古草群落	乔灌草–柏2	100°10′11.52″E	26°39′35.87″N	2660	重度	植树造林	6.28±0.14ab	55	28	18
干香柏–川滇野丁香–西南野古草群落	乔灌草–柏3	100°10′36.11″E	26°39′49.36″N	2584	重度	植树造林	6.17±0.14b	56	25	9
干香柏–小叶栒子–西南野古草群落	乔灌草–柏4	100°10′33.51″E	26°39′44.65″N	2570	重度	植树造林	6.29±0.07ab	55	25	10
干香柏–小叶鼠李–西南野古草群落	乔灌草-柏5	100°13′4.04″E	26°42′33.89″N	2544	重度	植树造林	5.62±0.15cd	65	35	8
云南松–西南野古草群落	纯松1	100°10′32.75″E	26°39′54.89″N	2637	轻度	封山育林	5.38±0.06d	8	52	30
云南松-西南野古草群落	纯松2	100°13′28.54″E	26°42′52.41″N	2600	轻度	封山育林	5.63±0.38cd	10	74	25
云南松–西南野古草群落	纯松3	100°18′45.49″E	27°01′26.42″N	2843	中度	植树造林	6.14±0.08ab	49	47	15
云南松–矮生胡枝子群落	纯松4	100°17′25.29″E	27°01′35.70″N	3078	轻度	植树造林	6.22±0.11ab	42	40	20
云南松–矮生胡枝子群落	纯松5	100°17′04.59″E	27°02′14.67″N	3057	轻度	植树造林	5.87±0.21bc	15	60	25
云南松–高山栎–云南草沙蚕群落	乔灌草–松1	100°16′57.52″E	27°02′11.38″N	3084	轻度	封山育林	5.39±0.05d	25	60	20
云南松–高山栎–云南草沙蚕群落	乔灌草–松2	100°16′57.29″E	27°02′9.92″N	3060	中度	封山育林	5.68±0.07d	42	48	20
云南松–帽斗栎–羊茅群落	乔灌草-松3	100°17′26.55″E	27°01′34.91″N	3080	轻度	封山育林	6.18±0.18b	20	60	18
云南松–帽斗栎–羊茅群落	乔灌草–松4	100°17′17.36″E	27°01′40.27″N	3070	中度	封山育林	5.77±0.05cd	41	42	18
云南松–柊树–西南野古草群落	乔灌草–松5	100°18′42.20″E	27°01′9.08″N	2821	中度	植树造林	6.02±0.10b	40	44	13
注：群落类型数字1~3与1~5表示在该群落类型内的每一个群落样点，与图1和图2中的数字相对应；“abcd”表示显著性差异（P<0.05）。 Note: The numbers 1 to 3 and 1 to 5 in community succession stages represent each community sample point within that succession stage and correspond to the numbers in Figure 1 and 2; “abcd” indicates significant differences (P < 0.05).

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