植物生态学报 ›› 2013, Vol. 37 ›› Issue (6): 542-550.DOI: 10.3724/SP.J.1258.2013.00055
收稿日期:
2013-01-04
接受日期:
2013-03-11
出版日期:
2013-01-04
发布日期:
2013-06-05
通讯作者:
杨洪晓
基金资助:
ZHANG Min1,PAN Yan-Xia2,YANG Hong-Xiao1,*()
Received:
2013-01-04
Accepted:
2013-03-11
Online:
2013-01-04
Published:
2013-06-05
Contact:
YANG Hong-Xiao
摘要:
沿海岸线分布的潮上带沙滩是受风暴潮影响的独特生境, 蕴含着独特的植物群落与植物资源。很多沙滩已受到不同程度的人为干扰。人们尚未了解潮上带沙滩植物群落的构建方式, 也忽视了各种人为干扰对它的影响。该文以几何级数模型、分割线段模型、重叠生态位模型和中性理论模型, 剖析了山东半岛潮上带3处典型沙草地的物种多度格局, 同时研究了滩涂养殖和旅游践踏两种最重要的干扰方式对潮上带沙滩植物群落的影响。对于这3处植被, 中性理论模型的拟合效果最好, 其次是几何级数模型, 而分割线段模型和重叠生态位模型的拟合效果不够理想。旅游践踏和沙滩养殖使潮上带沙草地的植被盖度降低, 不利于珍稀特有植物的生存。在潮上带沙滩上很多植物具有不少相似的特征, 导致它们在个体水平上的空间替代有强烈的随机性, 服从群落中性理论。然而, 它们的性状毕竟存在或多或少的差异, 这些差异难免对各种植物的资源分配或竞争能力产生微弱的影响, 经过很多年的积累, 这种影响必然在群落水平上逐渐显现出来, 以至于用几何级数模型也可拟合物种的多度格局。作者建议应控制人为干扰对潮上带沙滩珍稀植物的不利影响。
张敏,潘艳霞,杨洪晓. 山东半岛潮上带沙草地的物种多度格局及其对人为干扰的响应. 植物生态学报, 2013, 37(6): 542-550. DOI: 10.3724/SP.J.1258.2013.00055
ZHANG Min,PAN Yan-Xia,YANG Hong-Xiao. Species abundance patterns of supratidal sandy grassland along China’s Shandong Peninsula and their responses to human disturbances. Chinese Journal of Plant Ecology, 2013, 37(6): 542-550. DOI: 10.3724/SP.J.1258.2013.00055
物种或裸地 Species or bare land | 习性 Habit | 生活型 Life form | 高度 Height (cm) | 寿命 Lifetime (a) | 根型 Root form | 根深 Root depth (cm) | 近原生状态的物种频率 Species frequency in nearly original state (%) | 池塘堤坝的物种频率 Species frequency on pond walls (%) | 旅游践踏状态的物种频率 Species frequency under tourist trampling (%) |
---|---|---|---|---|---|---|---|---|---|
裸地 Bare land | 11.60 | 18.40 | 39.10 | ||||||
沙钻薹草 Carex kobomugi | CP | G | <50 | >2 | F | <50 | 49.41 | 41.96 | 3.33 |
粗毛鸭嘴草 Ischaemum bartatum | CP | G | <50 | >2 | F | <50 | 31.09 | 1.27 | 1.96 |
肾叶打碗花 Calystegia soldanella | CP | G | <50 | >2 | T | <50 | 18.22 | 27.16 | 11.67 |
砂引草 Messerschmidia sibirica | CP | G | <50 | >2 | T | <50 | 13.17 | 2.16 | 7.25 |
拟漆姑 Spergularia marina | CP | G | <50 | <2 | T | <50 | 12.97 | ||
滨麦 Leymus mollis | CP | G | <50 | >2 | F | <50 | 5.84 | 23.33 | |
矮生薹草 Carex pumila | CP | G | <50 | >2 | F | <50 | 3.47 | 6.47 | 13.73 |
沙滩黄芩 Scutellaria strigillosa | CP | G | <50 | >2 | T | <50 | 2.08 | ||
珊瑚菜 Glehnia littoralis | CP | G | <50 | >2 | T | >50 | 1.49 | ||
单叶蔓荆 Vitex trifolia var. simplicifolia | CP | W | <50 | >2 | T | >50 | 0.89 | 7.94 | |
沙苦荬菜 Chorisis repens | CP | G | <50 | >2 | T | <50 | 0.10 | 0.29 | 0.59 |
海滨香豌豆 Lathyrus maritimus | CP | G | <50 | >2 | T | <50 | 1.47 | ||
月见草 Oenothera erythrosepala | NTP | G | <50 | <2 | T | <50 | 2.77 | ||
独行菜 Lepidium apetalum | NTP | G | <50 | <2 | T | <50 | 2.57 | ||
猪毛菜 Salsola collina | NTP | G | <50 | <2 | T | <50 | 1.39 | 4.90 | 2.25 |
白茅 Imperata cylindrica | NTP | G | <50 | >2 | F | <50 | 0.99 | 2.94 | |
小蓬草 Conyza canadensis | NTP | G | >50 | <2 | T | <50 | 0.99 | ||
兴安胡枝子 Lespedeza davurica | NTP | W | <50 | >2 | T | <50 | 0.30 | ||
稗 Echinochloa crusgalli | NTP | G | >50 | <2 | F | <50 | 0.20 | 0.59 | 0.10 |
芦苇 Phragmites australis | NTP | G | >50 | >2 | F | <50 | 0.10 | 0.10 | |
白茎盐生草 Halogeton arachnoideus | NTP | G | <50 | <2 | T | <50 | 0.10 | ||
酸模 Rumex acetosa | NTP | G | >50 | >2 | T | <50 | 0.10 | ||
紫穗槐 Amorpha fruticosa | NTP | W | >50 | >2 | T | >50 | 0.10 | ||
狗牙根 Cynodon dactylon | NTP | G | <50 | >2 | F | <50 | 4.41 | 19.41 | |
香附子 Cyperus rotundus | NTP | G | <50 | >2 | F | <50 | 13.04 | ||
西伯利亚蓼 Polygonum sibiricum | NTP | G | <50 | >2 | T | <50 | 2.16 | ||
盐生草 Halogeton glomeratus | NTP | G | <50 | <2 | T | <50 | 0.69 | 1.37 | |
盐地碱蓬 Suaeda salsa | NTP | G | <50 | <2 | T | <50 | 0.29 | ||
苍耳 Xanthium sibiricum | NTP | G | >50 | <2 | T | <50 | 0.20 | ||
灰绿藜 Chenopodium glaucum | NTP | G | <50 | <2 | T | <50 | 0.20 | ||
酸模叶蓼 Polygonum lapathifolium | NTP | G | >50 | <2 | T | <50 | 0.20 | ||
紫苜蓿 Medicago sativa | NTP | G | >50 | >2 | T | <50 | 0.20 | ||
藜 Chenopodium album | NTP | G | <50 | <2 | T | <50 | 0.10 | 0.10 | |
巴天酸模 Rumex patientia | NTP | G | >50 | >2 | T | <50 | 0.10 | ||
刺沙蓬 Salsola ruthenica | NTP | G | >50 | <2 | T | <50 | 0.10 | ||
虫实 Corispermum spp. | NTP | G | <50 | <2 | T | <50 | 0.88 | ||
马齿苋 Portulaca oleracea | NTP | G | <50 | <2 | T | <50 | 0.10 |
表1 潮上带沙草地在3种条件下的物种组成
Table 1 Species composition of supratidal sandy grassland in three different conditions
物种或裸地 Species or bare land | 习性 Habit | 生活型 Life form | 高度 Height (cm) | 寿命 Lifetime (a) | 根型 Root form | 根深 Root depth (cm) | 近原生状态的物种频率 Species frequency in nearly original state (%) | 池塘堤坝的物种频率 Species frequency on pond walls (%) | 旅游践踏状态的物种频率 Species frequency under tourist trampling (%) |
---|---|---|---|---|---|---|---|---|---|
裸地 Bare land | 11.60 | 18.40 | 39.10 | ||||||
沙钻薹草 Carex kobomugi | CP | G | <50 | >2 | F | <50 | 49.41 | 41.96 | 3.33 |
粗毛鸭嘴草 Ischaemum bartatum | CP | G | <50 | >2 | F | <50 | 31.09 | 1.27 | 1.96 |
肾叶打碗花 Calystegia soldanella | CP | G | <50 | >2 | T | <50 | 18.22 | 27.16 | 11.67 |
砂引草 Messerschmidia sibirica | CP | G | <50 | >2 | T | <50 | 13.17 | 2.16 | 7.25 |
拟漆姑 Spergularia marina | CP | G | <50 | <2 | T | <50 | 12.97 | ||
滨麦 Leymus mollis | CP | G | <50 | >2 | F | <50 | 5.84 | 23.33 | |
矮生薹草 Carex pumila | CP | G | <50 | >2 | F | <50 | 3.47 | 6.47 | 13.73 |
沙滩黄芩 Scutellaria strigillosa | CP | G | <50 | >2 | T | <50 | 2.08 | ||
珊瑚菜 Glehnia littoralis | CP | G | <50 | >2 | T | >50 | 1.49 | ||
单叶蔓荆 Vitex trifolia var. simplicifolia | CP | W | <50 | >2 | T | >50 | 0.89 | 7.94 | |
沙苦荬菜 Chorisis repens | CP | G | <50 | >2 | T | <50 | 0.10 | 0.29 | 0.59 |
海滨香豌豆 Lathyrus maritimus | CP | G | <50 | >2 | T | <50 | 1.47 | ||
月见草 Oenothera erythrosepala | NTP | G | <50 | <2 | T | <50 | 2.77 | ||
独行菜 Lepidium apetalum | NTP | G | <50 | <2 | T | <50 | 2.57 | ||
猪毛菜 Salsola collina | NTP | G | <50 | <2 | T | <50 | 1.39 | 4.90 | 2.25 |
白茅 Imperata cylindrica | NTP | G | <50 | >2 | F | <50 | 0.99 | 2.94 | |
小蓬草 Conyza canadensis | NTP | G | >50 | <2 | T | <50 | 0.99 | ||
兴安胡枝子 Lespedeza davurica | NTP | W | <50 | >2 | T | <50 | 0.30 | ||
稗 Echinochloa crusgalli | NTP | G | >50 | <2 | F | <50 | 0.20 | 0.59 | 0.10 |
芦苇 Phragmites australis | NTP | G | >50 | >2 | F | <50 | 0.10 | 0.10 | |
白茎盐生草 Halogeton arachnoideus | NTP | G | <50 | <2 | T | <50 | 0.10 | ||
酸模 Rumex acetosa | NTP | G | >50 | >2 | T | <50 | 0.10 | ||
紫穗槐 Amorpha fruticosa | NTP | W | >50 | >2 | T | >50 | 0.10 | ||
狗牙根 Cynodon dactylon | NTP | G | <50 | >2 | F | <50 | 4.41 | 19.41 | |
香附子 Cyperus rotundus | NTP | G | <50 | >2 | F | <50 | 13.04 | ||
西伯利亚蓼 Polygonum sibiricum | NTP | G | <50 | >2 | T | <50 | 2.16 | ||
盐生草 Halogeton glomeratus | NTP | G | <50 | <2 | T | <50 | 0.69 | 1.37 | |
盐地碱蓬 Suaeda salsa | NTP | G | <50 | <2 | T | <50 | 0.29 | ||
苍耳 Xanthium sibiricum | NTP | G | >50 | <2 | T | <50 | 0.20 | ||
灰绿藜 Chenopodium glaucum | NTP | G | <50 | <2 | T | <50 | 0.20 | ||
酸模叶蓼 Polygonum lapathifolium | NTP | G | >50 | <2 | T | <50 | 0.20 | ||
紫苜蓿 Medicago sativa | NTP | G | >50 | >2 | T | <50 | 0.20 | ||
藜 Chenopodium album | NTP | G | <50 | <2 | T | <50 | 0.10 | 0.10 | |
巴天酸模 Rumex patientia | NTP | G | >50 | >2 | T | <50 | 0.10 | ||
刺沙蓬 Salsola ruthenica | NTP | G | >50 | <2 | T | <50 | 0.10 | ||
虫实 Corispermum spp. | NTP | G | <50 | <2 | T | <50 | 0.88 | ||
马齿苋 Portulaca oleracea | NTP | G | <50 | <2 | T | <50 | 0.10 |
模型 Model | 真实值与理论值比较 Comparison of real data with theoretical data | 近原生状态 Nearly original state | 池塘堤坝 Pond walls | 旅游条件 Tourist trampling |
---|---|---|---|---|
中性理论模型 Neutral theoretical model | 相对偏差 Relative deviation | 30.9% | 25.2% | 37.7% |
K-S检验显著性 K-S test significance | 0.871 | 1.000 | 0.632 | |
Pearson相关系数 Pearson correlation coefficient | 0.995 | 0.982 | 0.943 | |
AIC值 Aikake’s information criterion | 115.88 | 106.16 | 141.26 | |
几何级数模型 Geometric-series model | 相对偏差 Relative deviation | 34.3% | 31.9% | 45.2% |
K-S检验显著性 K-S test significance | 0.632 | 1.000 | 0.872 | |
Pearson相关系数 Pearson correlation coefficient | 0.980 | 0.956 | 0.864 | |
AIC值 Aikake’s information criterion | 154.04 | 123.69 | 157.94 | |
分割线段模型 Broken stick model | 相对偏差 Relative deviation | 71.0% | 64.7% | 65.7% |
K-S检验显著性 K-S test significance | 0.109 | 0.215 | 0.049 | |
Pearson相关系数 Pearson correlation coefficient | 0.943 | 0.957 | 0.963 | |
AIC值 Aikake’s information criterion | 184.13 | 129.63 | 140.87 | |
重叠生态位模型 Overlapping niche model | 相对偏差 Relative deviation | 85.8% | 77.0% | 76.0% |
K-S检验显著性 K-S test significance | 0.020 | 0.093 | 0.049 | |
Pearson相关系数 Pearson correlation coefficient | 0.827 | 0.874 | 0.898 | |
AIC值 Aikake’s information criterion | 196.44 | 139.99 | 156.35 |
表2 4种理论模型的拟合效果
Table 2 Fitting effects of four theoretical models
模型 Model | 真实值与理论值比较 Comparison of real data with theoretical data | 近原生状态 Nearly original state | 池塘堤坝 Pond walls | 旅游条件 Tourist trampling |
---|---|---|---|---|
中性理论模型 Neutral theoretical model | 相对偏差 Relative deviation | 30.9% | 25.2% | 37.7% |
K-S检验显著性 K-S test significance | 0.871 | 1.000 | 0.632 | |
Pearson相关系数 Pearson correlation coefficient | 0.995 | 0.982 | 0.943 | |
AIC值 Aikake’s information criterion | 115.88 | 106.16 | 141.26 | |
几何级数模型 Geometric-series model | 相对偏差 Relative deviation | 34.3% | 31.9% | 45.2% |
K-S检验显著性 K-S test significance | 0.632 | 1.000 | 0.872 | |
Pearson相关系数 Pearson correlation coefficient | 0.980 | 0.956 | 0.864 | |
AIC值 Aikake’s information criterion | 154.04 | 123.69 | 157.94 | |
分割线段模型 Broken stick model | 相对偏差 Relative deviation | 71.0% | 64.7% | 65.7% |
K-S检验显著性 K-S test significance | 0.109 | 0.215 | 0.049 | |
Pearson相关系数 Pearson correlation coefficient | 0.943 | 0.957 | 0.963 | |
AIC值 Aikake’s information criterion | 184.13 | 129.63 | 140.87 | |
重叠生态位模型 Overlapping niche model | 相对偏差 Relative deviation | 85.8% | 77.0% | 76.0% |
K-S检验显著性 K-S test significance | 0.020 | 0.093 | 0.049 | |
Pearson相关系数 Pearson correlation coefficient | 0.827 | 0.874 | 0.898 | |
AIC值 Aikake’s information criterion | 196.44 | 139.99 | 156.35 |
图1 潮上带沙草地的物种多度格局。把物种从常见到稀有排序, 以其序号作为横坐标。A, 近原生状态的潮上带沙草地, 位于胶南。B, 池塘堤坝的潮上带沙草地, 位于牟平。C, 遭受旅游践踏的潮上带沙草地, 位于蓬莱。
Fig. 1 Species-abundance patterns of supratidal sandy grassland. The horizontal axis refers to ranks of species in their more-to-less sorting. A, Supratidal sandy grassland in nearly original state in Jiaonan. B, Supratidal sandy grassland on pond walls in Muping. C, Supratidal sandy grassland suffering from tourist trampling in Penglai.
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