生物多样性 ›› 2014, Vol. 22 ›› Issue (2): 141-156. DOI: 10.3724/SP.J.1003.2014.13195
王斌, 黄俞淞, 李先琨*(), 向悟生, 丁涛, 黄甫昭, 陆树华, 韩文衡, 文淑均, 何兰军
收稿日期:
2013-08-30
接受日期:
2013-10-25
出版日期:
2014-03-20
发布日期:
2014-04-03
通讯作者:
李先琨
基金资助:
Bin Wang, Yusong Huang, Xiankun Li*(), Wusheng Xiang, Tao Ding, Fuzhao Huang, Shuhua Lu, Wenheng Han, Shujun Wen, Lanjun He
Received:
2013-08-30
Accepted:
2013-10-25
Online:
2014-03-20
Published:
2014-04-03
Contact:
Li Xiankun
摘要:
北热带喀斯特季节性雨林(northern tropical karst seasonal rain forest)是在我国热带北缘喀斯特地区分布的典型森林植被类型之一。由于富钙偏碱的地球化学背景及多样性的生境类型, 同时受季风气候影响, 该森林呈现群落结构多样、树种组成丰富、特有成分突出等特点。基于大型固定监测样地对该森林树种组成与空间分布进行的研究, 是探明该区域生物多样性形成与维持机制的基础。我们于2011年底建立了广西弄岗北热带喀斯特季节性雨林15 ha监测样地, 依照CTFS (Center for Tropical Forest Science)全球森林生物多样性监测规范, 定位并调查了样地内每一棵胸径≥1 cm的木本植物。结果表明: (1)样地内有监测树种223种, 隶属于56科157属; 独立个体总数为68,010株(含分枝为95,471株), 平均胸径为4.84 cm; (2)树种科、属的区系均以热带成分为主, 大戟科、马鞭草科、梧桐科等为优势科; (3)个体数最多的11个树种的个体数之和占到总个体数的51.64%, 前58个树种的占90.19%; 稀有种有75种, 占总树种数的33.63%; (4)群落结构稳定且更新良好, 主要优势种的径级结构均呈倒“J”形, 无明显断层; (5)树种分布在空间上表现出明显差异, 黄梨木(Boniodendron minus)等强耐旱型树种分布于山顶周围; 蚬木(Excentrodendron tonkinense)等树种分布于山坡中部; 对叶榕(Ficus hispida)等喜湿耐荫型树种分布于山谷周围; (6)胸径>20 cm的个体较多分布在山坡中下部, 极少分布在山顶周围; 胸径10–20 cm的个体较均匀分布于整个样地; 分枝和萌枝较多分布在山顶周围; (7)种–面积散点图在2–7 ha的取样面积下分化形成两条曲线, 表明了树种数量组成在空间上具有强烈异质性。研究初步认为: 强烈生境异质性及独特地质背景可能是影响该喀斯特森林物种组成及空间分布的重要因素。
王斌, 黄俞淞, 李先琨, 向悟生, 丁涛, 黄甫昭, 陆树华, 韩文衡, 文淑均, 何兰军 (2014) 弄岗北热带喀斯特季节性雨林15 ha监测样地的树种组成与空间分布. 生物多样性, 22, 141-156. DOI: 10.3724/SP.J.1003.2014.13195.
Bin Wang,Yusong Huang,Xiankun Li,Wusheng Xiang,Tao Ding,Fuzhao Huang,Shuhua Lu,Wenheng Han,Shujun Wen,Lanjun He (2014) Species composition and spatial distribution of a 15 ha northern tropical karst seasonal rain forest dynamics study plot in Nonggang, Guangxi, southern China. Biodiversity Science, 22, 141-156. DOI: 10.3724/SP.J.1003.2014.13195.
图1 弄岗自然保护区1971–2000年的月平均降雨量、气温、风速的变化
Fig. 1 Monthly variations of rainfall, air temperature and wind speed between 1971 and 2000 in the Nonggang Nature Reserve
图2 弄岗样地地形图及其在中国的地理位置。根据植被及土壤分布规律, 弄岗样地初步可分3种生境类型: (1)海拔>320 m的山顶类型; (2)海拔位于240–320 m之间的山坡类型; (3)海拔<240 m的洼地类型。
Fig. 2 The location and contour map of the 15-ha Nonggang forest plot, China. According to the vegetation and soil distribution, the Nonggang forest plot was basically divided into three habitat types: (1) Mountain peak type at the altitude > 320 m; (2) Mountain slope type at the altitude between 240 and 320 m; (3) Valley bottom type at the altitude < 240 m.
分布区类型 Areal types | 科数 No. of families | 科的比例 % | 属数 No. of genera | 属的比例 % |
---|---|---|---|---|
1. 世界广布 Cosmopolitan | 12 | – | 2 | – |
2. 泛热带 Pantropic | 31 | 70.45 | 36 | 23.23 |
3. 热带亚洲及热带美洲间断 Tropical Asia and South Tropical America disjuncted | 5 | 11.36 | 6 | 3.87 |
4. 旧世界热带 Old World Tropic | 2 | 4.54 | 23 | 14.84 |
5. 热带亚洲至热带大洋洲 Tropical Asia to Tropical Australasia Oceania | 0 | 0 | 18 | 11.61 |
6. 热带亚洲至热带非洲 Tropical Asia to Tropical Africa | 0 | 0 | 9 | 5.81 |
7. 热带亚洲 Tropical Asia | 1 | 2.27 | 45 | 29.03 |
8. 北温带 North Temperate | 4 | 9.09 | 5 | 3.23 |
9. 东亚和北美洲间断 East Asia and North America disjuncted | 1 | 2.27 | 7 | 4.52 |
11. 温带亚洲 Temperate Asia | 0 | 0 | 1 | 0.65 |
12. 地中海区、西亚至中亚 Mediterranean, West Asia to Central Asia | 0 | 0 | 1 | 0.65 |
14. 东亚 East Asia | 0 | 0 | 4 | 2.58 |
合计 Total | 56 | 100 | 157 | 100 |
表1 弄岗森林样地木本植物区系类型
Table 1 The areal-types of woody plants in the Nonggang forest plot, Guangxi
分布区类型 Areal types | 科数 No. of families | 科的比例 % | 属数 No. of genera | 属的比例 % |
---|---|---|---|---|
1. 世界广布 Cosmopolitan | 12 | – | 2 | – |
2. 泛热带 Pantropic | 31 | 70.45 | 36 | 23.23 |
3. 热带亚洲及热带美洲间断 Tropical Asia and South Tropical America disjuncted | 5 | 11.36 | 6 | 3.87 |
4. 旧世界热带 Old World Tropic | 2 | 4.54 | 23 | 14.84 |
5. 热带亚洲至热带大洋洲 Tropical Asia to Tropical Australasia Oceania | 0 | 0 | 18 | 11.61 |
6. 热带亚洲至热带非洲 Tropical Asia to Tropical Africa | 0 | 0 | 9 | 5.81 |
7. 热带亚洲 Tropical Asia | 1 | 2.27 | 45 | 29.03 |
8. 北温带 North Temperate | 4 | 9.09 | 5 | 3.23 |
9. 东亚和北美洲间断 East Asia and North America disjuncted | 1 | 2.27 | 7 | 4.52 |
11. 温带亚洲 Temperate Asia | 0 | 0 | 1 | 0.65 |
12. 地中海区、西亚至中亚 Mediterranean, West Asia to Central Asia | 0 | 0 | 1 | 0.65 |
14. 东亚 East Asia | 0 | 0 | 4 | 2.58 |
合计 Total | 56 | 100 | 157 | 100 |
科名 Family | 分布区类型 Areal types | 属数 No. of genera | 树种数 No. of species | 个体数 No. of stems | 断面积之和 Basal area (m2) | 重要值IV |
---|---|---|---|---|---|---|
大戟科 Euphorbiaceae | 泛热带 Pantropic | 21 | 36 | 23,181 | 61.6 | 19.78 |
马鞭草科 Verbenaceae | 热带亚洲及热带南美间断 Tropical Asia and North America disjuncted | 4 | 9 | 6,985 | 48.1 | 8.67 |
梧桐科 Sterculiaceae | 泛热带 Pantropic | 4 | 5 | 11,514 | 37.81 | 8.62 |
桑科 Moraceae | 世界广布 Cosmopolitan | 6 | 18 | 5,667 | 16.42 | 6.36 |
椴树科 Tiliaceae | 泛热带 Pantropic | 3 | 3 | 4,007 | 27.39 | 4.63 |
茜草科 Rubiaceae | 世界广布 Cosmopolitan | 9 | 11 | 2,578 | 7.81 | 3.35 |
楝科 Meliaceae | 世界广布 Cosmopolitan | 8 | 9 | 3,397 | 6.82 | 3.24 |
番荔枝科 Annonaceae | 泛热带 Pantropic | 6 | 7 | 3,531 | 2.54 | 2.55 |
无患子科 Sapindaceae | 泛热带 Pantropic | 6 | 6 | 1,791 | 9.86 | 2.53 |
蝶形花科 Fabaceae | 世界广布 Cosmopolitan | 4 | 4 | 323 | 17.22 | 2.46 |
柿树科 Ebenaceae | 泛热带 Pantropic | 1 | 4 | 3,394 | 5.61 | 2.36 |
紫葳科 Bignoniaceae | 泛热带 Pantropic | 4 | 6 | 764 | 8.47 | 2.03 |
表2 弄岗森林样地重要值排名前12位的科
Table 2 Top 12 families with the highest importance values in the Nonggang forest plot, Guangxi
科名 Family | 分布区类型 Areal types | 属数 No. of genera | 树种数 No. of species | 个体数 No. of stems | 断面积之和 Basal area (m2) | 重要值IV |
---|---|---|---|---|---|---|
大戟科 Euphorbiaceae | 泛热带 Pantropic | 21 | 36 | 23,181 | 61.6 | 19.78 |
马鞭草科 Verbenaceae | 热带亚洲及热带南美间断 Tropical Asia and North America disjuncted | 4 | 9 | 6,985 | 48.1 | 8.67 |
梧桐科 Sterculiaceae | 泛热带 Pantropic | 4 | 5 | 11,514 | 37.81 | 8.62 |
桑科 Moraceae | 世界广布 Cosmopolitan | 6 | 18 | 5,667 | 16.42 | 6.36 |
椴树科 Tiliaceae | 泛热带 Pantropic | 3 | 3 | 4,007 | 27.39 | 4.63 |
茜草科 Rubiaceae | 世界广布 Cosmopolitan | 9 | 11 | 2,578 | 7.81 | 3.35 |
楝科 Meliaceae | 世界广布 Cosmopolitan | 8 | 9 | 3,397 | 6.82 | 3.24 |
番荔枝科 Annonaceae | 泛热带 Pantropic | 6 | 7 | 3,531 | 2.54 | 2.55 |
无患子科 Sapindaceae | 泛热带 Pantropic | 6 | 6 | 1,791 | 9.86 | 2.53 |
蝶形花科 Fabaceae | 世界广布 Cosmopolitan | 4 | 4 | 323 | 17.22 | 2.46 |
柿树科 Ebenaceae | 泛热带 Pantropic | 1 | 4 | 3,394 | 5.61 | 2.36 |
紫葳科 Bignoniaceae | 泛热带 Pantropic | 4 | 6 | 764 | 8.47 | 2.03 |
树种名 Species | 科名 Families | 个体数 No. of stems | 分枝数 No. of ramifications | 平均断面积Mean basal area (cm2) | 相对频度Relative frequency | 重要值IV | 偏好生境Preferred habitats |
---|---|---|---|---|---|---|---|
闭花木 Cleistanthus sumatranus | Euphorbiaceae | 9,977 | 2,005 | 30.13 | 1.50 | 9.05 | 中上坡 H-2 |
苹婆 Sterculia monosperma | Sterculiaceae | 6,328 | 2,738 | 37.76 | 2.79 | 7.47 | 中下坡 H-4 |
广西牡荆 Vitex kwangsiensis | Verbenaceae | 2,470 | 2,933 | 87.75 | 2.63 | 6.86 | 中下坡 H-4 |
蚬木 Excentrodendron tonkinense | Tiliaceae | 1,502 | 369 | 76.87 | 1.82 | 2.79 | 中上坡 H-2 |
海南椴 Diplodiscus trichosperma | Tiliaceae | 1,126 | 856 | 65.30 | 1.76 | 2.44 | 中上坡 H-2 |
劲直刺桐 Erythrina stricta | Fabaceae | 316 | 2 | 541.58 | 1.29 | 2.32 | 谷底 H-5 |
海南大风子 Hydnocarpus hainanensis | Flacourtiaceae | 2,260 | 333 | 14.21 | 2.41 | 2.28 | 中下坡 H-4 |
日本五月茶 Antidesma japonicum | Euphorbiaceae | 2,535 | 373 | 10.48 | 1.91 | 2.18 | 谷底 H-5 |
对叶榕 Ficus hispida | Moraceae | 2,989 | 188 | 14.11 | 0.68 | 2.14 | 谷底 H-5 |
金丝李 Garcinia paucinervis | Guttiferae | 1,684 | 258 | 21.96 | 2.60 | 2.12 | 中上坡 H-2 |
茎花山柚 Champereia manillana | Opiliaceae | 1,340 | 138 | 22.76 | 2.39 | 1.79 | 中坡 H-3 |
假玉桂 Celtis timorensis | Ulmaceae | 1,122 | 27 | 42.65 | 2.11 | 1.74 | 中下坡 H-4 |
细叶谷木 Memecylon scutellatum | Melastomataceae | 1,221 | 974 | 21.60 | 2.00 | 1.74 | 山顶 H-1 |
截裂翅子树 Pterospermum truncatolobatum | Sterculiaceae | 1,604 | 721 | 13.98 | 1.75 | 1.69 | 中坡 H-3 |
山榄叶柿 Diospyros siderophylla | Ebenaceae | 1,592 | 531 | 17.83 | 1.30 | 1.59 | 山顶 H-1 |
三角车 Rinorea bengalensis | Violaceae | 2,178 | 476 | 9.38 | 0.77 | 1.57 | 中坡 H-3 |
黄梨木 Boniodendron minus | Sapindaceae | 694 | 406 | 75.20 | 1.15 | 1.55 | 山顶 H-1 |
割舌树 Walsura robusta | Meliaceae | 1,315 | 246 | 18.69 | 1.67 | 1.49 | 中坡 H-3 |
肥牛树 Cephalomappa sinensis | Euphorbiaceae | 917 | 919 | 33.66 | 0.78 | 1.33 | 中上坡 H-2 |
网脉核果木 Drypetes perreticulata | Euphorbiaceae | 631 | 99 | 60.84 | 1.47 | 1.24 | 中坡 H-3 |
其他 Other | – | 24,209 | 12,869 | – | 65.22 | 44.62 | – |
总计 Total | – | 68,010 | 27,461 | – | 100 | 100 | – |
表3 弄岗森林样地重要值排名前20位树种的组成及偏好生境
Table 3 Composition and preferred habitats of top 20 species with the highest important values in the Nonggang forest plot, Guangxi
树种名 Species | 科名 Families | 个体数 No. of stems | 分枝数 No. of ramifications | 平均断面积Mean basal area (cm2) | 相对频度Relative frequency | 重要值IV | 偏好生境Preferred habitats |
---|---|---|---|---|---|---|---|
闭花木 Cleistanthus sumatranus | Euphorbiaceae | 9,977 | 2,005 | 30.13 | 1.50 | 9.05 | 中上坡 H-2 |
苹婆 Sterculia monosperma | Sterculiaceae | 6,328 | 2,738 | 37.76 | 2.79 | 7.47 | 中下坡 H-4 |
广西牡荆 Vitex kwangsiensis | Verbenaceae | 2,470 | 2,933 | 87.75 | 2.63 | 6.86 | 中下坡 H-4 |
蚬木 Excentrodendron tonkinense | Tiliaceae | 1,502 | 369 | 76.87 | 1.82 | 2.79 | 中上坡 H-2 |
海南椴 Diplodiscus trichosperma | Tiliaceae | 1,126 | 856 | 65.30 | 1.76 | 2.44 | 中上坡 H-2 |
劲直刺桐 Erythrina stricta | Fabaceae | 316 | 2 | 541.58 | 1.29 | 2.32 | 谷底 H-5 |
海南大风子 Hydnocarpus hainanensis | Flacourtiaceae | 2,260 | 333 | 14.21 | 2.41 | 2.28 | 中下坡 H-4 |
日本五月茶 Antidesma japonicum | Euphorbiaceae | 2,535 | 373 | 10.48 | 1.91 | 2.18 | 谷底 H-5 |
对叶榕 Ficus hispida | Moraceae | 2,989 | 188 | 14.11 | 0.68 | 2.14 | 谷底 H-5 |
金丝李 Garcinia paucinervis | Guttiferae | 1,684 | 258 | 21.96 | 2.60 | 2.12 | 中上坡 H-2 |
茎花山柚 Champereia manillana | Opiliaceae | 1,340 | 138 | 22.76 | 2.39 | 1.79 | 中坡 H-3 |
假玉桂 Celtis timorensis | Ulmaceae | 1,122 | 27 | 42.65 | 2.11 | 1.74 | 中下坡 H-4 |
细叶谷木 Memecylon scutellatum | Melastomataceae | 1,221 | 974 | 21.60 | 2.00 | 1.74 | 山顶 H-1 |
截裂翅子树 Pterospermum truncatolobatum | Sterculiaceae | 1,604 | 721 | 13.98 | 1.75 | 1.69 | 中坡 H-3 |
山榄叶柿 Diospyros siderophylla | Ebenaceae | 1,592 | 531 | 17.83 | 1.30 | 1.59 | 山顶 H-1 |
三角车 Rinorea bengalensis | Violaceae | 2,178 | 476 | 9.38 | 0.77 | 1.57 | 中坡 H-3 |
黄梨木 Boniodendron minus | Sapindaceae | 694 | 406 | 75.20 | 1.15 | 1.55 | 山顶 H-1 |
割舌树 Walsura robusta | Meliaceae | 1,315 | 246 | 18.69 | 1.67 | 1.49 | 中坡 H-3 |
肥牛树 Cephalomappa sinensis | Euphorbiaceae | 917 | 919 | 33.66 | 0.78 | 1.33 | 中上坡 H-2 |
网脉核果木 Drypetes perreticulata | Euphorbiaceae | 631 | 99 | 60.84 | 1.47 | 1.24 | 中坡 H-3 |
其他 Other | – | 24,209 | 12,869 | – | 65.22 | 44.62 | – |
总计 Total | – | 68,010 | 27,461 | – | 100 | 100 | – |
图9 弄岗样地所有分枝和萌枝在3个不同径级的空间分布图
Fig. 9 Spatial distribution maps of tree ramifications and sprouts at three different DBH classes in the Nonggang forest plot
样地名称 Forest plot | 地理坐标 Coordinates | 面积 Size (ha) | 科 Family | 属 Genus | 种 Species | 个体数 Stem (株/ha) | 胸高断面积 BA (m2/ha) | 稀有种 Rare (%) |
---|---|---|---|---|---|---|---|---|
1 长白山–阔叶红松林 | 42.38° N, 128.08° E | 25 | 18 | 32 | 52 | 1,556 | 43.23 | 34.60 |
2 天童山–中亚热带常绿阔叶林 | 29.81° N, 121.77° E | 20 | 50 | 91 | 153 | 4,730 | 38.98 | 36.20 |
3 古田山–中亚热带常绿阔叶林 | 29.25° N, 118.12° E | 24 | 49 | 104 | 159 | 5,863 | 36.90 | 37.10 |
4 鼎湖山–南亚热带常绿阔叶林 | 23.10° N, 112.32° E | 20 | 56 | 119 | 210 | 3,581 | 30.17 | 52.38 |
5 弄岗–北热带喀斯特季节性雨林 | 22.43° N, 106.95° E | 15 | 56 | 157 | 223 | 4,534 | 22.08 | 33.63 |
6 垦丁–热带高位珊瑚礁季雨林 | 21.95° N, 120.82° E | 10 | 34 | 74 | 95 | 3,909 | 45.15 | 28.42 |
7 西双版纳–热带雨林 | 21.61° N, 101.57° E | 20 | 70 | 213 | 468 | 4,792 | 42.34 | 49.14 |
表4 不同植被类型森林监测样地的地理坐标、面积以及树种组成(郝占庆等, 2008; 兰国玉等, 2008; 叶万辉等, 2008; 祝燕等, 2008; Wu et al., 2011; 杨庆松等, 2011)
Table 4 Location, size and species composition of stem-mapping forest plots with different vegetation types
样地名称 Forest plot | 地理坐标 Coordinates | 面积 Size (ha) | 科 Family | 属 Genus | 种 Species | 个体数 Stem (株/ha) | 胸高断面积 BA (m2/ha) | 稀有种 Rare (%) |
---|---|---|---|---|---|---|---|---|
1 长白山–阔叶红松林 | 42.38° N, 128.08° E | 25 | 18 | 32 | 52 | 1,556 | 43.23 | 34.60 |
2 天童山–中亚热带常绿阔叶林 | 29.81° N, 121.77° E | 20 | 50 | 91 | 153 | 4,730 | 38.98 | 36.20 |
3 古田山–中亚热带常绿阔叶林 | 29.25° N, 118.12° E | 24 | 49 | 104 | 159 | 5,863 | 36.90 | 37.10 |
4 鼎湖山–南亚热带常绿阔叶林 | 23.10° N, 112.32° E | 20 | 56 | 119 | 210 | 3,581 | 30.17 | 52.38 |
5 弄岗–北热带喀斯特季节性雨林 | 22.43° N, 106.95° E | 15 | 56 | 157 | 223 | 4,534 | 22.08 | 33.63 |
6 垦丁–热带高位珊瑚礁季雨林 | 21.95° N, 120.82° E | 10 | 34 | 74 | 95 | 3,909 | 45.15 | 28.42 |
7 西双版纳–热带雨林 | 21.61° N, 101.57° E | 20 | 70 | 213 | 468 | 4,792 | 42.34 | 49.14 |
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