长江口盐沼优势蟹类的生境选择与生态系统工程师效应
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摘要
河口盐沼是高产的但非常脆弱的、易受干扰的生态系统,同时也是蟹类生物多样性最高的区域之一。蟹类作为生态系统工程师,对湿地生态系统的物质循环和能量流动具有深刻影响。蟹类的物种组成和洞穴分布决定了蟹类生态功能的范围和强度,而外来物种入侵则可能深刻改变了蟹类的群落结构和生态系统功能。本论文通过野外调查、野外受控实验、室内食性实验等一系列实验方法,研究了长江口盐沼优势蟹类的生境选择与生态系统工程师效应。主要结果如下:
1)为了认识长江河口盐沼优势蟹无齿相手蟹的生境选择,在野外比较了互花米草、芦苇群落和裸地之间该蟹的多度和生物量差异。结果表明,互花米草生境中无齿相手蟹的多度和生物量显著高于芦苇和裸地。这是由于互花米草生境中的土壤含水量和植物群落特征显著不同于芦苇和裸地,互花米草群落更适合无齿相手蟹的栖息。摄食喜好实验结果表明,无齿相手蟹摄食互花米草的量是芦苇的两倍以上。互花米草通过向无齿相手蟹提供适口的食物和温和的环境条件,为其提供了适宜的生活环境。
2)植物凋落物的存在可能对生境具有深刻影响,从而影响蟹类的分布。为了确定凋落物对蟹类生境选择的影响,通过野外操作实验探讨了凋落物密度和凋落物状态对蟹类分布的影响。凋落物密度对无齿相手蟹种群和雄性的生物量具有显著影响,适中的凋落物密度适合蟹类的生存。互花米草凋落物对雄性无齿相手蟹生物量具有显著影响,移除凋落物的生境中蟹类分布最多。在春季,对照和凋落物全移除的样地中天津厚蟹雌性蟹体重显著高于具有中等密度凋落物的样地,而在其他月份不同凋落物密度处理之间没有显著差异。春季的雌性天津厚蟹体重在站立凋落物内比倒伏的生境更小,而在其他月份则没有显著差异。天津厚蟹的种群参数,除了性比以外,在芦苇和互花米草之间都没有显著差异,但植物群落却对无齿相手蟹种群产生了深刻的影响。
3)通过野外实验研究了盐沼中潮沟剖面环境异质性对蟹洞特征的影响,并比较了潮沟和非潮沟生境间蟹洞分布的差异。结果表明,在潮沟剖面上从底部到平滩,土壤含水量逐渐下降,含水量变化也引起了土壤其他物理、化学性质的变化。在斜坡上蟹洞密度高但是洞穴较小,在边沿和平滩洞穴较大而密度较低。这些洞穴分布的差异很可能是由蟹类的分带引起的。潮沟剖面上土壤性质呈现出从水生到陆地生境的过渡,这决定了蟹类在潮沟剖面上的分带。细粒土壤通常含有较多的粘滞性颗粒,所以也更稳定,在坡面洞穴开口总面积与土壤颗粒大小之间呈负相关关系。相对于非潮沟生境,潮沟内小洞穴的密度更高,但大洞穴的密度较低,而且潮沟内环境和生物组分的变异也更大。所以,潮沟剖面的环境异质性能满足不同蟹类的多样需求,并为蟹类提供了重要的生态交错区。
4)为了认识不同生境类型对蟹洞形态的影响,通过原位洞穴浇塑法研究了洞穴形态在芦苇、互花米草和裸地间的差异。植物群落通过改变根系分布、土壤性质和其他的环境因素,显著改变了蟹洞的形态。相对于植物群落,裸地的土壤含水量较低,但容重较高。芦苇和互花米草群落内的洞穴分枝多,弯曲度高,但洞穴深度、长度和体积较小。与芦苇相比,互花米草的高度较小,根状茎生物量更低,但是密度、细根生物量更高,所以互花米草群落中洞穴深度、长度、体积都较小,开口直径和弯曲度都比芦苇群落内洞穴要大。互花米草老种群的上层土壤容重较低,但含水量较高,而新种群具有更大的高度、地上生物量,较低的密度和根系生物量。互花米草新种群中的蟹洞比老种群中具有更大的深度、长度、宽度和体积,但是更低弯曲度。进一步的分析表明,土壤含水量、容重、植物根状茎和细根、地上生物量对蟹洞形态起决定性的作用。
5)蟹类掘穴对盐沼生态系统的物质循环和能量流动具有深刻影响;为了定量掘穴蟹类对盐沼土壤的生态系统工程师效应,通过野外实验研究了蟹类掘穴对土壤、碳、氮的垂直运输。蟹类掘穴行为引起了土壤含水量、碳、氮的增加和土壤容重的降低。蟹类对土壤和营养的周转率随着洞穴大小的增加而增加。植物(芦苇和互花米草)显著增加了土壤的营养浓度,但阻碍了蟹类对物质的垂直运输,限制了营养的循环和重新利用。蟹类挖掘的土壤量远高于沉积到蟹洞内的土壤量,导致了土壤向地面的净输出。芦苇、互花米草群落和裸地上蟹类净输出到地面的土壤量分别是171.7,109.5,375.0 g·m~(-2).d~(-1),三种生境中蟹类活动周转一次土壤的时间分别为2.89,4.07和1.83 yr。蟹类很显然是重要的生态系统工程师,在几年时间内能完全混合表层和深层土壤,加速碎屑分解和植物对营养的利用。
Coastal salt marshes are one of the most productive ecosystems but are fragile and sensitive to environmental changes. The salt marshes have a great diversity of burrowing crabs. As physical ecosystem engineers, crabs play important roles in controlling material cycles and energy flows in salt marsh ecosystems. Through a series of experimental methods, including field investigations, field controlled experiments and laboratory experiments, this study aimed to evaluate vertical movements of crabs' burrowing processes on soil, carbon and nitrogen in salt marsh, and analyze the effects of exotic plant invasions on crab distributions, burrow morphologies and ecological functions. The major findings are summarized as follows:
Abundance and biomass of Sesarma dehaani in Spartina marsh were significantly greater than those in Phragmites marsh and mudflats. Soil water content and plant community characteristics of Spartina marsh also significantly differed from those of Phragmites marsh and mudflats. Moreover, analysis of feeding preference showed that S. dehaani consumed Spartina more than twice as much Phragmites. Therefore, Spartina provided compatible habitats for native crab S. dehaani through offering suitable food source and moderate environmental conditions.
Plant litter had significant effects on some of crab community parameters. Spartina litter affected male S. dehaani biomass, and Spartina marsh with removal of litter had the highest crab density. In spring, body weight of female Helice tridens tientsinensis was the lowest in Spartina marsh with medium litter density; and it was lower in Spartina marsh with standing litter than with lodging litter. However, plant litter had no effects body weight of females in other seasons. Except for sex ratio, population parameters of H. tientsinensis were not significantly different between P. australis and S. alterniflora marshes, but those of S. dehaani had great defference between the marshes.
Sediment properties showed a gradual transition from hydrophytic to terrestrial environments on creek sections. On the slope, the burrows were smaller in size and burrow density was higher, while at the edge and on the flat, the burrows were larger in size, and the burrow density was lower. The density of small crab burrows was greater, but the density of large burrows and total burrow opening area were lower in tidal creeks than in non-creek habitats. The environmental heterogeneities on creek sections met the requirements of different crab species, and hence the creeks of tidal salt marshes provided important ecotones for crabs.
Plant communities significantly changed crab burrow morphologies through changing root distribution pattern, soil properties and other environmental conditions during the course of plant growth, plant invasion and plant colonization of mudflats in salt marshes. Burrows in Phragmites and Spartina marshes had higher branch number and sinuosity degree, but burrow depth, and volume compared to mudflats. Burrows in Spartina marshes had lower burrow depth and burrow volume, but higher opening diameter, sinuosity degree compared to Phragmites marshes. Burrows in young Spartina marshes had higher burrow depth and volume, but lower sinuosity degree than those in old marshes.
Crab burrowing increased soil water content, carbon and nitrogen and decreased soil bulk density; and the turnover of soil and nutrients by crabs increased with burrow size. Plants (Phragmites and Spartina) significantly enhanced soil nutrient concentrations, but obstructed the vertical movement of materials through crab burrowing, limiting nutrient cycling and reuse. The burrowing behavior of crabs led to a net transport of soil to the surface. Burrowing crabs are clearly important ecosystem engineers that can entirely mix surface and deeper soils over a period of one-four years and accelerate litter decomposition and efficient reuse of nutrients by plants.
1)为了认识长江河口盐沼优势蟹无齿相手蟹的生境选择,在野外比较了互花米草、芦苇群落和裸地之间该蟹的多度和生物量差异。结果表明,互花米草生境中无齿相手蟹的多度和生物量显著高于芦苇和裸地。这是由于互花米草生境中的土壤含水量和植物群落特征显著不同于芦苇和裸地,互花米草群落更适合无齿相手蟹的栖息。摄食喜好实验结果表明,无齿相手蟹摄食互花米草的量是芦苇的两倍以上。互花米草通过向无齿相手蟹提供适口的食物和温和的环境条件,为其提供了适宜的生活环境。
2)植物凋落物的存在可能对生境具有深刻影响,从而影响蟹类的分布。为了确定凋落物对蟹类生境选择的影响,通过野外操作实验探讨了凋落物密度和凋落物状态对蟹类分布的影响。凋落物密度对无齿相手蟹种群和雄性的生物量具有显著影响,适中的凋落物密度适合蟹类的生存。互花米草凋落物对雄性无齿相手蟹生物量具有显著影响,移除凋落物的生境中蟹类分布最多。在春季,对照和凋落物全移除的样地中天津厚蟹雌性蟹体重显著高于具有中等密度凋落物的样地,而在其他月份不同凋落物密度处理之间没有显著差异。春季的雌性天津厚蟹体重在站立凋落物内比倒伏的生境更小,而在其他月份则没有显著差异。天津厚蟹的种群参数,除了性比以外,在芦苇和互花米草之间都没有显著差异,但植物群落却对无齿相手蟹种群产生了深刻的影响。
3)通过野外实验研究了盐沼中潮沟剖面环境异质性对蟹洞特征的影响,并比较了潮沟和非潮沟生境间蟹洞分布的差异。结果表明,在潮沟剖面上从底部到平滩,土壤含水量逐渐下降,含水量变化也引起了土壤其他物理、化学性质的变化。在斜坡上蟹洞密度高但是洞穴较小,在边沿和平滩洞穴较大而密度较低。这些洞穴分布的差异很可能是由蟹类的分带引起的。潮沟剖面上土壤性质呈现出从水生到陆地生境的过渡,这决定了蟹类在潮沟剖面上的分带。细粒土壤通常含有较多的粘滞性颗粒,所以也更稳定,在坡面洞穴开口总面积与土壤颗粒大小之间呈负相关关系。相对于非潮沟生境,潮沟内小洞穴的密度更高,但大洞穴的密度较低,而且潮沟内环境和生物组分的变异也更大。所以,潮沟剖面的环境异质性能满足不同蟹类的多样需求,并为蟹类提供了重要的生态交错区。
4)为了认识不同生境类型对蟹洞形态的影响,通过原位洞穴浇塑法研究了洞穴形态在芦苇、互花米草和裸地间的差异。植物群落通过改变根系分布、土壤性质和其他的环境因素,显著改变了蟹洞的形态。相对于植物群落,裸地的土壤含水量较低,但容重较高。芦苇和互花米草群落内的洞穴分枝多,弯曲度高,但洞穴深度、长度和体积较小。与芦苇相比,互花米草的高度较小,根状茎生物量更低,但是密度、细根生物量更高,所以互花米草群落中洞穴深度、长度、体积都较小,开口直径和弯曲度都比芦苇群落内洞穴要大。互花米草老种群的上层土壤容重较低,但含水量较高,而新种群具有更大的高度、地上生物量,较低的密度和根系生物量。互花米草新种群中的蟹洞比老种群中具有更大的深度、长度、宽度和体积,但是更低弯曲度。进一步的分析表明,土壤含水量、容重、植物根状茎和细根、地上生物量对蟹洞形态起决定性的作用。
5)蟹类掘穴对盐沼生态系统的物质循环和能量流动具有深刻影响;为了定量掘穴蟹类对盐沼土壤的生态系统工程师效应,通过野外实验研究了蟹类掘穴对土壤、碳、氮的垂直运输。蟹类掘穴行为引起了土壤含水量、碳、氮的增加和土壤容重的降低。蟹类对土壤和营养的周转率随着洞穴大小的增加而增加。植物(芦苇和互花米草)显著增加了土壤的营养浓度,但阻碍了蟹类对物质的垂直运输,限制了营养的循环和重新利用。蟹类挖掘的土壤量远高于沉积到蟹洞内的土壤量,导致了土壤向地面的净输出。芦苇、互花米草群落和裸地上蟹类净输出到地面的土壤量分别是171.7,109.5,375.0 g·m~(-2).d~(-1),三种生境中蟹类活动周转一次土壤的时间分别为2.89,4.07和1.83 yr。蟹类很显然是重要的生态系统工程师,在几年时间内能完全混合表层和深层土壤,加速碎屑分解和植物对营养的利用。
Coastal salt marshes are one of the most productive ecosystems but are fragile and sensitive to environmental changes. The salt marshes have a great diversity of burrowing crabs. As physical ecosystem engineers, crabs play important roles in controlling material cycles and energy flows in salt marsh ecosystems. Through a series of experimental methods, including field investigations, field controlled experiments and laboratory experiments, this study aimed to evaluate vertical movements of crabs' burrowing processes on soil, carbon and nitrogen in salt marsh, and analyze the effects of exotic plant invasions on crab distributions, burrow morphologies and ecological functions. The major findings are summarized as follows:
Abundance and biomass of Sesarma dehaani in Spartina marsh were significantly greater than those in Phragmites marsh and mudflats. Soil water content and plant community characteristics of Spartina marsh also significantly differed from those of Phragmites marsh and mudflats. Moreover, analysis of feeding preference showed that S. dehaani consumed Spartina more than twice as much Phragmites. Therefore, Spartina provided compatible habitats for native crab S. dehaani through offering suitable food source and moderate environmental conditions.
Plant litter had significant effects on some of crab community parameters. Spartina litter affected male S. dehaani biomass, and Spartina marsh with removal of litter had the highest crab density. In spring, body weight of female Helice tridens tientsinensis was the lowest in Spartina marsh with medium litter density; and it was lower in Spartina marsh with standing litter than with lodging litter. However, plant litter had no effects body weight of females in other seasons. Except for sex ratio, population parameters of H. tientsinensis were not significantly different between P. australis and S. alterniflora marshes, but those of S. dehaani had great defference between the marshes.
Sediment properties showed a gradual transition from hydrophytic to terrestrial environments on creek sections. On the slope, the burrows were smaller in size and burrow density was higher, while at the edge and on the flat, the burrows were larger in size, and the burrow density was lower. The density of small crab burrows was greater, but the density of large burrows and total burrow opening area were lower in tidal creeks than in non-creek habitats. The environmental heterogeneities on creek sections met the requirements of different crab species, and hence the creeks of tidal salt marshes provided important ecotones for crabs.
Plant communities significantly changed crab burrow morphologies through changing root distribution pattern, soil properties and other environmental conditions during the course of plant growth, plant invasion and plant colonization of mudflats in salt marshes. Burrows in Phragmites and Spartina marshes had higher branch number and sinuosity degree, but burrow depth, and volume compared to mudflats. Burrows in Spartina marshes had lower burrow depth and burrow volume, but higher opening diameter, sinuosity degree compared to Phragmites marshes. Burrows in young Spartina marshes had higher burrow depth and volume, but lower sinuosity degree than those in old marshes.
Crab burrowing increased soil water content, carbon and nitrogen and decreased soil bulk density; and the turnover of soil and nutrients by crabs increased with burrow size. Plants (Phragmites and Spartina) significantly enhanced soil nutrient concentrations, but obstructed the vertical movement of materials through crab burrowing, limiting nutrient cycling and reuse. The burrowing behavior of crabs led to a net transport of soil to the surface. Burrowing crabs are clearly important ecosystem engineers that can entirely mix surface and deeper soils over a period of one-four years and accelerate litter decomposition and efficient reuse of nutrients by plants.
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