中国西南喀斯特瀑布苔藓植物的生态学研究
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摘要
喀斯特瀑布(Karst waterfalls)是独特的生态系统之一,具有其特殊的生态学、地质学和环境学意义。中国西南是世界上喀斯特瀑布分布数量最集中、类型最多和经济价值最大的区域之一。瀑布生物喀斯特沉积景观是一批世界自然遗产、国家级风景名胜区及自然保护区的重要组成成分,如贵州的黄果树瀑布群、四川的九寨沟瀑布群和黄龙瀑布群、云南的罗平九龙瀑布群和广西的德天瀑布等,具有重要研究价值。
本文以中国西南典型喀斯特瀑布为代表,采用生态学、植物地理学、喀斯特学和沉积学多学科交叉的理论和方法,于2004年10月至2007年8月,对86个喀斯特瀑布苔藓植物进行了较系统的野外调查,行程约3.6万公里,共采集样品和标本2100份;对黔中6种苔藓生长速度进行了为期两年的野外定点观测;利用A Analyst 800原子吸收光谱仪和AF-640原子荧光光谱仪对8种苔藓和水体中7种金属元素含量进行了分析;比较了中国西南和西欧喀斯特瀑布中分布的苔藓群落组成特征。
结果如下:
1)依据水源特征并结合喀斯特地貌特点,提出了一个新的喀斯特瀑布分类系统。将该区喀斯特瀑布划分为2大类型、4个亚类型和12个小类型,并揭示了其地理分布规律性。
2)该区域瀑布苔藓植物共有34科92属248种(含变种亚种)。依据各物种分布频度,结果表明罕见种占54.03%、稀见种占32.66%、偶见种占6.86%、频见种占4.43%和常见种占2.02%。该区苔藓地理成分为热带亚洲成分占20.97%、泛热带成分占1.21%、热带亚洲至热带非洲成分占1.21%、东亚成分占19.35%、北温带成分占20.57%、东亚和北美洲际间断成分占1.61%、旧世界温带成分占1.21%、温带亚洲成分占4.03%、中国特有占14.5%和世界广布种占15.32%。
3)依据对钙的适应性能力的高低,可将苔藓植物划分钙华苔藓、喜钙苔藓和广幅苔藓等3种适钙生态类群,报道了中国钙华苔藓38种。发现瀑布区苔藓营养繁殖普遍,孢子体罕见,雌雄异株的种类占总数的81.45%等特点。苔藓适应瀑布沉积环境,植物体形态上相应出现了绿色生长区、半钙化区、钙化区和石化区等分化。植物体钙化和石化的结果,增强了苔藓适应瀑布富钙、高速沉积和高速冲击力的激流环境的能力。
4)黔中6种苔藓植物配子体地上部分年均生长速度是:大叶风尾藓是1.283cm/yr、狄氏石灰藓是2.072cm/yr、威氏石灰藓是2.35cm/yr、钩喙净口藓是1.76cm/yr、橙色净口藓是0.883cm/yr和牛角藓是1.56cm/yr。苔藓植物的生长速度因种类不同及生长季节有差异。
5)在喀斯特瀑布中生长的苔藓植物体内金属元素含量普遍比较高,含量的顺序与水体中的顺序基本一致。不同物种植物体对金属元素的吸收、富集却有较大的差异。根据8种苔藓对瀑布水体中不同金属元素的富集作用的差异可划分为4组:大叶石灰藓和东亚泽藓对Zn和Ca超强烈富集;橙色净口藓和湿原藓对Zn和Cu超强烈富集:狄氏石灰藓和牛角藓对Zn超强烈富集;仰叶藓和大叶风尾藓对Cu、Pb和Zn等三种元素超强烈富集。
6)中国西南18个喀斯特瀑布中,共有30种苔藓植物群落,由12科27属61种组成;而西欧18个喀斯特瀑布中,仅有28种苔藓植物群落,由15科24属33种组成。尽管两地代表性的群落略有差异,但总体上拥有许多相似的特征,如生活型和生态分布类型等。
7)根据水流和地貌特点,划分中国西南喀斯特瀑布苔藓沉积物为4大生态沉积类型;新报道了16种苔藓喀斯特生态沉积小类型。
8)提出了喀斯特瀑布沉积生境中急需保护的20种苔藓物种建议名单。
Karst waterfalls are one of the special ecosystems that interest ecologists, geologists and environmental scientists. China is one of countries in the world with thousands of karst wa- terfalls of different types and economic importance. The landscapes of karst waterfalls are important for a lot of world heritage sites, national natural reserves and national scenic spots in southwestern China, for example, Huangguoshu Waterfalls in Guizhou, Jiuzhaigou Waterfalls and Huanglong Waterfalls in Sichuan, Nine-Dragon Waterfalls in Yunnan and Detian Waterfalls in Guangxi. All of them are important for biodiversity conservation, tourism and basic science research.
From October 2004 to August 2007, many surveys were conducted in the karst area of southwestern China. About 2100 samples and specimens were collected from 86 active karst waterfalls. The growth rates of six aquatic mosses were measured at eight karst water- falls in the field from August 2005 to August 2007. By means of Atomic Absorption Spectrometry (A Analyst 800) and Atomic Fluorescence Spectroscopy (AF-640), contents of seven metal elements of eight mosses and water samples were analysed from eight karst waterfalls in Guizhou Province. To understand the general features of aquatic bryophyte communities of karst waterfalls, we made a comparison of bryophyte communities of 36 karst waterfalls from southwestern China and western Europe.
Major results are summarised as follows:
1) Based on a series of investigations for 86 karst waterfalls in southwestern China, a classification system of karst waterfalls was proposed. According to the characteristics of karst water sources and karst landforms, karst waterfalls were divided into two types, and subdivided into four subtypes and further into twelve small types. Distribution and features of karst waterfalls were discussed.
2) The bryoflora of karst waterfalls consisted of 248 species (including varieties and subspecies) in the southwestern China, which belong to 92 genera and 34 families. Five occurrence frequencies of bryophytes in karst waterfalls were distinguished: very rare species (134 taxa, 54.03%), rare species (81 taxa, 32.66%), frequent species (11 taxa, 4.43%), occasional species (17 taxa, 6.86%) and common species (5 taxa, 2.02%). The phytogeo-graphical elements of the bryoflora included Tropical Asian (20.97%), Pantropical (1.21 %), Tropic Asian to Tropic African (1.21%), East Asian (19.35%), North Temperate (20.57%), East Asian and North America disjuncted ( 1.61% ), Old World Temperate (1.21%), Temperate Asian elements (4.03%), Endemics to China(14.5%) and Cosmopolitans (15.32%).
3) According to the differences of bryophytes in ecological adaptation to calcite ions, three ecological groups were identified: travertine bryophytes (38 taxa, 15.32 % ), calciphilous bryophytes(98 taxa, 39.52 %) and eurytopic bryophytes (112 taxa, 45.16%). Vegetative propagations of bryophytes were common while sporophytes with capules were very rare. The reproductive systems consisted of five types, dioecious taxa accounted for 81.45%. Interestingly, we found that there were four different areas in shoots of mosses that might be associated with ecological adaptation, i.e., green growth area, half-calcification area, calcification area and petrification area. Calcification and petrification of shoots allow bryophytes to be successful competitors in harsh environments characterised by fast water flow, rich calcite and rapid deposition.
4) Growth rates were obtained for six mosses from eight karst waterfalls every month in Guizhou province from August 2005 to August 2007. The estimated rates per annum were as follows: Fissidens grandifrons 1.283 cm, Hydrogonium dixonianum 2.072 cm, H, williamsii 2.35 cm, Gymnostomum recurvirostre 1.76 cm, G aurantiacum 0.883 cm, Cra-toneuron filicinum 1.56 cm. The results were discussed with reference to karst depo si- tion, water chemistry and water flow rate. Seasonal patterns of growth of six moss species were investigated in field, and significant differences in relative growth rate were observed between species.
5) The results showed that the element contents in plants were higher, and that the element contents in plants were related to those in water. Moreover, the capacities to absorb and enrich elements varied among bryophytes. We found that Hydrogonium majusculum and Philonotis turneriana had strong ability to enrich Zn and Ca, Gymnostomum aurantiac- um and Calliergon cordifolium Zn and Cu, Hydrogonium dixoniana and Cratoneuron filicinum Zn, and Reimersia inconspicua and Fissidens grandifrons Cu, Pb and Zn. Therefore, our results suggest that these bryophytes may have ecological tolerance of different metal elements in water from karst waterfall in the area, hence can be used as indicators of metal contamination.
6) Comparative studies of characteristics of bryophyte communities were conducted between 18 karst waterfalls in southwestern China and 18 karst waterfalls in western Europe. We found that 30 aquatic bryophyte communities consisted of 61 taxa belonging to 27 genera and 12 families in China, whereas 28 aquatic bryophyte communities consisted of only 33 taxa belonging to 24 genera and 15 families in Western Europe. Although there were some differences in typical bryophyte communities, we found common features of southwestern China and western Europe, including life-forms and ecological distribution types of bryophyte communities.
7) A new classification for bryoliths associated with karst waterfalls was proposed. According to the features of environmental factors and bryophyte communities, four ecological types (stream waterfall deposition, river waterfall deposition, dam deposition and cave deposition ) and 16 subtypes of biokarst depositions associated with bryophytes could be distinguished.
8) From the viewpoint of biodiversity conservation of bryophytes in the karst area, 20 bryophyte taxa were suggested to be priority species for biodiversity conservation at karst waterfalls in southwstern China, i.e., Fissidens grandifrons Brid., Hydrogonium dixonianum Chen, H. majusculum (C. Muell.) Chen, H. gracileutum (Mitt.) Chen, H. consguireum (Thwait et Mitt.) Hilp., H. setschwanicum (Broth.)Chen, H. williamsii Chen, Gymnostomum recurvirostrum Hedw., G subridulum (Broth) Chen, G aurantiacum(Mitt.)Jaeg., G aeruginosum Sm, Anoectangium clarum Mitt., Cratoneuron filicinum (Hedw.) Spruce., Palustriella commutata (Hedw.) Ochyra, P. falcata (Brid.) Hedenas, Philonotis calcarea (B.S.G) Schimp., P. turneriana (Schwaegr.) Mitt., Bryum pseudotriquetrum (Hedw.) Gaerth, Barbula tophacea (Brid.) Mitt. and Cyathodium cavernarum Kunze.
本文以中国西南典型喀斯特瀑布为代表,采用生态学、植物地理学、喀斯特学和沉积学多学科交叉的理论和方法,于2004年10月至2007年8月,对86个喀斯特瀑布苔藓植物进行了较系统的野外调查,行程约3.6万公里,共采集样品和标本2100份;对黔中6种苔藓生长速度进行了为期两年的野外定点观测;利用A Analyst 800原子吸收光谱仪和AF-640原子荧光光谱仪对8种苔藓和水体中7种金属元素含量进行了分析;比较了中国西南和西欧喀斯特瀑布中分布的苔藓群落组成特征。
结果如下:
1)依据水源特征并结合喀斯特地貌特点,提出了一个新的喀斯特瀑布分类系统。将该区喀斯特瀑布划分为2大类型、4个亚类型和12个小类型,并揭示了其地理分布规律性。
2)该区域瀑布苔藓植物共有34科92属248种(含变种亚种)。依据各物种分布频度,结果表明罕见种占54.03%、稀见种占32.66%、偶见种占6.86%、频见种占4.43%和常见种占2.02%。该区苔藓地理成分为热带亚洲成分占20.97%、泛热带成分占1.21%、热带亚洲至热带非洲成分占1.21%、东亚成分占19.35%、北温带成分占20.57%、东亚和北美洲际间断成分占1.61%、旧世界温带成分占1.21%、温带亚洲成分占4.03%、中国特有占14.5%和世界广布种占15.32%。
3)依据对钙的适应性能力的高低,可将苔藓植物划分钙华苔藓、喜钙苔藓和广幅苔藓等3种适钙生态类群,报道了中国钙华苔藓38种。发现瀑布区苔藓营养繁殖普遍,孢子体罕见,雌雄异株的种类占总数的81.45%等特点。苔藓适应瀑布沉积环境,植物体形态上相应出现了绿色生长区、半钙化区、钙化区和石化区等分化。植物体钙化和石化的结果,增强了苔藓适应瀑布富钙、高速沉积和高速冲击力的激流环境的能力。
4)黔中6种苔藓植物配子体地上部分年均生长速度是:大叶风尾藓是1.283cm/yr、狄氏石灰藓是2.072cm/yr、威氏石灰藓是2.35cm/yr、钩喙净口藓是1.76cm/yr、橙色净口藓是0.883cm/yr和牛角藓是1.56cm/yr。苔藓植物的生长速度因种类不同及生长季节有差异。
5)在喀斯特瀑布中生长的苔藓植物体内金属元素含量普遍比较高,含量的顺序与水体中的顺序基本一致。不同物种植物体对金属元素的吸收、富集却有较大的差异。根据8种苔藓对瀑布水体中不同金属元素的富集作用的差异可划分为4组:大叶石灰藓和东亚泽藓对Zn和Ca超强烈富集;橙色净口藓和湿原藓对Zn和Cu超强烈富集:狄氏石灰藓和牛角藓对Zn超强烈富集;仰叶藓和大叶风尾藓对Cu、Pb和Zn等三种元素超强烈富集。
6)中国西南18个喀斯特瀑布中,共有30种苔藓植物群落,由12科27属61种组成;而西欧18个喀斯特瀑布中,仅有28种苔藓植物群落,由15科24属33种组成。尽管两地代表性的群落略有差异,但总体上拥有许多相似的特征,如生活型和生态分布类型等。
7)根据水流和地貌特点,划分中国西南喀斯特瀑布苔藓沉积物为4大生态沉积类型;新报道了16种苔藓喀斯特生态沉积小类型。
8)提出了喀斯特瀑布沉积生境中急需保护的20种苔藓物种建议名单。
Karst waterfalls are one of the special ecosystems that interest ecologists, geologists and environmental scientists. China is one of countries in the world with thousands of karst wa- terfalls of different types and economic importance. The landscapes of karst waterfalls are important for a lot of world heritage sites, national natural reserves and national scenic spots in southwestern China, for example, Huangguoshu Waterfalls in Guizhou, Jiuzhaigou Waterfalls and Huanglong Waterfalls in Sichuan, Nine-Dragon Waterfalls in Yunnan and Detian Waterfalls in Guangxi. All of them are important for biodiversity conservation, tourism and basic science research.
From October 2004 to August 2007, many surveys were conducted in the karst area of southwestern China. About 2100 samples and specimens were collected from 86 active karst waterfalls. The growth rates of six aquatic mosses were measured at eight karst water- falls in the field from August 2005 to August 2007. By means of Atomic Absorption Spectrometry (A Analyst 800) and Atomic Fluorescence Spectroscopy (AF-640), contents of seven metal elements of eight mosses and water samples were analysed from eight karst waterfalls in Guizhou Province. To understand the general features of aquatic bryophyte communities of karst waterfalls, we made a comparison of bryophyte communities of 36 karst waterfalls from southwestern China and western Europe.
Major results are summarised as follows:
1) Based on a series of investigations for 86 karst waterfalls in southwestern China, a classification system of karst waterfalls was proposed. According to the characteristics of karst water sources and karst landforms, karst waterfalls were divided into two types, and subdivided into four subtypes and further into twelve small types. Distribution and features of karst waterfalls were discussed.
2) The bryoflora of karst waterfalls consisted of 248 species (including varieties and subspecies) in the southwestern China, which belong to 92 genera and 34 families. Five occurrence frequencies of bryophytes in karst waterfalls were distinguished: very rare species (134 taxa, 54.03%), rare species (81 taxa, 32.66%), frequent species (11 taxa, 4.43%), occasional species (17 taxa, 6.86%) and common species (5 taxa, 2.02%). The phytogeo-graphical elements of the bryoflora included Tropical Asian (20.97%), Pantropical (1.21 %), Tropic Asian to Tropic African (1.21%), East Asian (19.35%), North Temperate (20.57%), East Asian and North America disjuncted ( 1.61% ), Old World Temperate (1.21%), Temperate Asian elements (4.03%), Endemics to China(14.5%) and Cosmopolitans (15.32%).
3) According to the differences of bryophytes in ecological adaptation to calcite ions, three ecological groups were identified: travertine bryophytes (38 taxa, 15.32 % ), calciphilous bryophytes(98 taxa, 39.52 %) and eurytopic bryophytes (112 taxa, 45.16%). Vegetative propagations of bryophytes were common while sporophytes with capules were very rare. The reproductive systems consisted of five types, dioecious taxa accounted for 81.45%. Interestingly, we found that there were four different areas in shoots of mosses that might be associated with ecological adaptation, i.e., green growth area, half-calcification area, calcification area and petrification area. Calcification and petrification of shoots allow bryophytes to be successful competitors in harsh environments characterised by fast water flow, rich calcite and rapid deposition.
4) Growth rates were obtained for six mosses from eight karst waterfalls every month in Guizhou province from August 2005 to August 2007. The estimated rates per annum were as follows: Fissidens grandifrons 1.283 cm, Hydrogonium dixonianum 2.072 cm, H, williamsii 2.35 cm, Gymnostomum recurvirostre 1.76 cm, G aurantiacum 0.883 cm, Cra-toneuron filicinum 1.56 cm. The results were discussed with reference to karst depo si- tion, water chemistry and water flow rate. Seasonal patterns of growth of six moss species were investigated in field, and significant differences in relative growth rate were observed between species.
5) The results showed that the element contents in plants were higher, and that the element contents in plants were related to those in water. Moreover, the capacities to absorb and enrich elements varied among bryophytes. We found that Hydrogonium majusculum and Philonotis turneriana had strong ability to enrich Zn and Ca, Gymnostomum aurantiac- um and Calliergon cordifolium Zn and Cu, Hydrogonium dixoniana and Cratoneuron filicinum Zn, and Reimersia inconspicua and Fissidens grandifrons Cu, Pb and Zn. Therefore, our results suggest that these bryophytes may have ecological tolerance of different metal elements in water from karst waterfall in the area, hence can be used as indicators of metal contamination.
6) Comparative studies of characteristics of bryophyte communities were conducted between 18 karst waterfalls in southwestern China and 18 karst waterfalls in western Europe. We found that 30 aquatic bryophyte communities consisted of 61 taxa belonging to 27 genera and 12 families in China, whereas 28 aquatic bryophyte communities consisted of only 33 taxa belonging to 24 genera and 15 families in Western Europe. Although there were some differences in typical bryophyte communities, we found common features of southwestern China and western Europe, including life-forms and ecological distribution types of bryophyte communities.
7) A new classification for bryoliths associated with karst waterfalls was proposed. According to the features of environmental factors and bryophyte communities, four ecological types (stream waterfall deposition, river waterfall deposition, dam deposition and cave deposition ) and 16 subtypes of biokarst depositions associated with bryophytes could be distinguished.
8) From the viewpoint of biodiversity conservation of bryophytes in the karst area, 20 bryophyte taxa were suggested to be priority species for biodiversity conservation at karst waterfalls in southwstern China, i.e., Fissidens grandifrons Brid., Hydrogonium dixonianum Chen, H. majusculum (C. Muell.) Chen, H. gracileutum (Mitt.) Chen, H. consguireum (Thwait et Mitt.) Hilp., H. setschwanicum (Broth.)Chen, H. williamsii Chen, Gymnostomum recurvirostrum Hedw., G subridulum (Broth) Chen, G aurantiacum(Mitt.)Jaeg., G aeruginosum Sm, Anoectangium clarum Mitt., Cratoneuron filicinum (Hedw.) Spruce., Palustriella commutata (Hedw.) Ochyra, P. falcata (Brid.) Hedenas, Philonotis calcarea (B.S.G) Schimp., P. turneriana (Schwaegr.) Mitt., Bryum pseudotriquetrum (Hedw.) Gaerth, Barbula tophacea (Brid.) Mitt. and Cyathodium cavernarum Kunze.
引文
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