巨桉—台湾桤木混合凋落物分解特征及其土壤动物群落动态
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摘要
凋落物分解是森林地力维持的基础,而土壤动物是凋落物分解的积极参与者,并受到凋落物种类、质量和分解阶段的影响。因此,深入研究人工林凋落物分解过程及其土壤动物群落动态,可为人工林培育和可持续经营与管理提供重要的科学依据。巨桉(Eucalyptus grandis)是南方工业原料林的主要造林树种之一,而台湾桤木(Alnus formosana)是非豆科固氮的常绿速生树种,为了最大限度地避免桉树人工林地力衰退、生物多样性下降等生态学问题,研究人员开展了巨桉与台湾桤木两树种的混交试验。现在的问题是：如何控制桉树与混交树种之间的比例才能有效维持混交林的地力和保育生物多样性?树种混交是否影响到凋落物的分解过程?是否影响到凋落物中的土壤动物群落动态?迄今尚缺乏深入研究。以四川省乐山市苏稽镇巨桉-台湾桤木人工林为研究对象,采用凋落物分解袋法,研究了5种不同比例(TⅠ：巨桉10g,台湾桤木0g；TⅡ：巨桉7g,台湾桤木3g；TⅢ：巨桉5g,台湾桤木5g；TⅣ：巨桉3g,台湾桤木7g；TV：巨桉0g,台湾桤木10g)的混合凋落物在3种不同孔径(6目、30目和260目)尼龙袋(20cm×20cm)的分解特征以及土壤动物群落动态,以期充分认识巨桉与台湾桤木混合对凋落物分解速率、养分动态和土壤动物群落结构的影响,为构建合理可行的人工林模式提供理论依据。
(1)桉-桤不同混合比例凋落物中优势类群为弹尾目(Collembola)和蜱螨目(Acariformes)。常见类群为大型土壤动物后孔寡毛目(Opithopora)。五种不同混合比例的凋落物中,大型土壤动物的分布是TV最多,TⅠ最少,混合凋落物介于二者之间,且随桤木比例的增加呈增多的趋势；而中小型土壤动物在TⅠ中最多,而TV最少。啮虫目(Psocoptera)在TV中的数量远远高于其它四种凋落物,后孔寡毛目在混合凋落物中数量较高。弹尾目、蝎线纲(Pauropoda)更喜生活在TⅠ凋落物中。综上表明在巨桉中加入台湾桤木可改变土壤动物的群落结构。
(2)6目凋落袋中土壤动物的个体数是TⅠ>TⅡ>TⅢ>TⅣ>TV,大型土壤动物的个体数是TⅢ>TⅣ>TⅤ>TⅡ>TⅠ,中小型土壤动物个体数是TⅠ>TⅡ>TⅢ>TⅣ>TⅤ,大型土壤动物类群数在TV最多,混合凋落物次之,TⅠ最少。30目凋落袋中土壤动物个体数TⅠ>TⅡ>TV>TⅣ>TⅢ。260目凋落袋中,土壤动物的个体数是Ⅳ>TⅢ>TⅡ>TⅠ>TⅤ。五种不同比例混合凋落物中的土壤动物个体总数在6目中差异极其显著,在30目和260目差异性不显著。中小型土壤动物个体数量是260目>30目>6目,且在30目与6目中差异显著。6目中中小型土壤动物的个体数远高于大型土壤动物。土壤动物类群数是30目>6目>260目,五种不同比例混合凋落物中的土壤动物类群数在260目差异不显著,在30目差异显著,在6目中差异极其显著。表明相对纯桉而言,桉-桤混和可提高凋落物中大型土壤动物的个体数、类群数,而对中小型土壤动物的个体数和类群数并无影响。
(3)6目凋落袋内凋落物的分解速率为TV>TⅣ>TⅢ>TⅡ>TⅠ,TⅠ凋落物的快速分解期在8月,TⅡ和TⅢ凋落物的快速分解期在7月,而TⅣ和TV凋落物的快速分解期在5月。分解前期TⅡ由于桤木凋落物的加入,大大提高了整个凋落物的分解速度,而TⅣ因桉树凋落物的存在,分解速率有所降低。在2009年10月前,260目和30目凋落物袋中,TⅠ凋落物的分解速率快于TV。近自然状态下(6目中),分解前期(0-45d)由于桤木的加入TⅡ凋落物的分解速度有所提高,也即TⅡ(巨桉7g,台湾桤木3g)混合比例可促进凋落物的分解。
(4)背景土壤动物包括湿生中小型土壤动物、干生中小型土壤动物和大型土壤动物。湿生中小型土壤动物中线虫纲(Nematoda)为优势类群,轮虫纲(Rotatoria)和线蚓科(Enchytraeidae)为常见类群。干生中小型土壤动物优势类群有弹尾目和蜱螨目。大型土壤动物优势类群有后孔寡毛目、等翅目(Ⅰsoptera)、鞘翅目(Coleoptera)和膜翅目(Hymenoptera)。在五种类型的人工林中,湿生中小型土壤动物、干生中小型土壤动物以及大型土壤动物的个体数在TI人工林下中均较多,而在混交林和TV林下较少。土壤动物在不同土层个体数差异较大,尤其是大型土壤动物,表聚现象明显。
(5)不同比例桉-桤混合初始凋落物间氮差异极其显著,钙、镁差异显著,碳、磷、钾差异不显著。巨桉、台湾桤木两种凋落物间C：N比差异极其显著。在分解过程中,6目凋落袋中桉-桤不同混合比例凋落物各元素的含量都有所降低。C、K元素含量在TⅠ凋落物中下降最慢,Mg元素含量在TⅠ中下降最快。N、P元素含量在TV凋落物中下降最慢,C、Ca元素含量在TV中下降最快。30目凋落袋中N元素含量在TV凋落物中,C、Ca在TⅠ凋落物中下降缓慢,P、Mg元素含量在TⅠ凋落物中下降较快。260目凋落袋中N、Mg元素含量在TⅠ凋落物中下降迅速,在TV中下降缓慢。6目中凋落物的损失率与N元素、Ca元素的含量呈极其显著正相关,与P元素的含量呈显著正相关,与C含量呈负相关,但相关性不显著。
(6)260目凋落袋中凋落物的养分含量总体上高于30目，也即260目凋落物袋中凋落物的养分释放速率慢于30目,而260目的土壤动物个体数高于30目,土壤动物的类群数低于30目。表明不同的土壤动物类群有利于促进凋落物中养分元素的释放。大型土壤动物对各元素释放作用的大小依次为：Mg>P>K>N>Ca>C,Mg元素释放作用在TV凋落物最高,达92.45%,对C元素释放作用在TⅢ凋落物中最低,仅32.05%,对N元素的释放作用在TⅠ凋落物中最高。
研究表明：不同比例桉-桤混合可提高凋落物中大型土壤动物的个体数与类群数,加速凋落物的分解,促进养分的释放与循环。因此在桉树纯林中混栽桤木(比例为7：3),有利于加速人工林内的物质循环,提高土壤地力,促进植物生长。
Litter decomposition plays an essential role in maintaining soil fertility in the forest. Soil fauna which affected by the type and quality of litter and the stage of decomposition are active participants during the decomposition of litter. Therefore, the research on dynamics of soil fauna community during litter decay can provide important scientific basis for the cultivation and the sustainable management of plantation. Eucalyptus(Eucalyptus grandis) is one of main afforestation tree species in the industrial raw material forest in the south, and alder (Alnus formosana) is a nonleguminous and evergreen tree species with the characteristic of fast growing and nitrogen-fixing. In order to maximum extently avoid the ecological problems, such as soil fertility and biodiversity decline, the litter mixed experiment of Eucalyptus grandis and Alnus formosana was conducted. The corresponding problems were put up:(1) How do control the ratio of eucalyptus litter weight to alder litter weight to maintain productivity and biodiversity in the mixed forest?(2) Will different tree species mixture affect the litter decomposition process?(3) Will different tree species mixture change dynamitic of soil fauna community? All those are lack of corresponding research. Decomposition characteristic and dynamic of soil fauna in different ratio mixed litters (TⅠ:eucalyptus10g, alder Og; TⅡ:eucalyptus7g, alder3g; TⅢ:eucalyptus5g, alder5g; TⅣ:eucalyptus3g, alder7g; TⅤ:eucalyptus Og, alder10g) were investigated with the method of litterbags in Suji town, Leshan city, Sichuan province. The aim was to explore the effect brought by mixture of eucalyptus and alder on decomposition rate, dynamics of nutrient and soil fauna community structure and provide the support of science and technology for the building of reasonable and feasible plantation.
(1) The dominant groups were Collembola and Acariformes which belong to meso and microfauna. The common group was Opithopora which belongs to macrofauna in different ratio mixed litter. Compared with TI litter, the macrofauna preferred to live in TV litter. The individual numbers of macrofauna in mixed litter were between TV litter and TI litter. The individual numbers of macrofauna increases with quantity of the alder added. Compared with TV litter, the meso and microfauna preferred to live in TI litter. The individual numbers of Psocoptera reached highest in TV litter and Opithopora in mixed litter. Collembola and Pauropoda preferred living in TI litter. All these suggested: the community structure of soil fauna would change when alder was added in Eucalyptus.
(2) The individual numbers of soil fauna in6-mesh litterbags were TI>TII>TIII>TIV>TV, and those of macrofauna were TIII>TIV>TV>TII>TI and meso and microfauna were TⅠ>TⅡ>TⅢ>TⅣ>TⅤ. The group numbers of macrofauna were maximum in TV litter and minimum in TI litter. The individual numbers of soil fauna in30-mesh litterbags were TI>TII> TV>TIV>TIII and in260-mesh litterbags were IV> TⅢ>TⅡ>TⅠ>TⅤ. The total individuals of soil fauna showed extremely significant difference in6-mesh litterbags, no significant difference in260-mesh litterbags and30-mesh litterbags among five different ratio mixed litter. The individual numbers of meso and micro fauna were260-mesh>30-mesh>6-mesh and there was significant difference between30-mesh and6-mesh litterbags. The individual numbers of meso and microfauna were more than those of marofauna in6-mesh litterbags. The group numbers were30-mesh>6-mesh>260-mesh and there was significant difference in30-mesh litterbags, extremely significant difference in6-mesh litterbags and no significant difference in260-mesh litterbags among five different ratio mixed litter. All these showed the mixture of eucalyptus and alder could increase the individuals of soil macrofauna, whereas no effect on the individuals and groups of meso and microfauna, compared with the eucalyptus litter.
(3) The decomposition rate was TV>TIV>TIII>TII>TI in6-mesh litterbags, and the fast decomposition stage of TI was in August, TⅡ and TⅢ in July and TIV and TV in May. The decomposition rate of TⅡ litter increased greatly due to the addition of alder in early stage, and the decomposition rate of TIV litter declined because of the eucalyptus added. Before October2009, the decomposition rate of TI litter was faster than that of TⅤ litter in260-and30-mesh litterbags. Under near natural state (6-mesh), the ratio of TⅡ could accelerate the decay of litter.
(4) Background soil fauna included damp living meso and micro fauna, dry living meso and micro fauna and-macrofauna. The-dominant-group was Nematoda and the common groups were Rotatoria and Enchytraeidae among damp living meso and micro fauna. The dominant groups were Collembola and Acariformes among dry living meso-, micro fauna. The dominant groups were Opithopora, Isoptera, Coleoptera and Hymenoptera among macrofauna. There were more individuals in soil under TI plantation than mixed and TV plantation. The soil fauna, especially macrofauna, showed significant difference among different soil layer and more soil fauna happened in the top soil.
(5) There was extremely significant difference in initial N, significant difference in initial Ca and Mg, and no significant difference in C, P and K among different ratio mixed litter. The ratio of C:N showed extremely significant difference between eucalyptus and alder. The elements concentration of different ratio mixed littler declined in6-mesh litterbags during decomposition. The elements of C, K declined slowly and Mg fast in TI litter. The elements of N, P declined slowly and C, Ca fast in TV litter. The element of N, C and Ca declined slowly in TV, TI litter respectively, P, Mg declined fast in TI litter in30-mesh litterbags. N and Mg declined fast in TI litter and slowly in TV litter in260-mesh litterbags. Mass loss rate was extremely significant positively correlated to the concentration of N, Ca, significant correlated to concentration of P, negatively correlate to the content of C, but there was no obvious difference.
(6) The concentration of elements in260-mesh litterbags was higher than in30-mesh litterbags, that is to say, the nutrient release rate of litter in260-mesh litterbags was slower than in30-mesh litterbags, however, the individual numbers of soil fauna were higher in260-mesh litterbags than in-30-mesh litterbags and the group numbers were on the contrary. Different groups of soil fauna could stimulate the release of nutrient elements. The release rate of nutrient elements which was attribute to the soil macrofauna showed the order:Mg>P>K>N>Ca>C. Action of macrofauna to Mg was highest, achieving92.5%, however to C was lowest (32.05%), to N was highest in TI litter.
All the research suggested that the mixture of Eucalyptus grandis and Alnus formosana could increase the numbers of individual and group of macrofauna, accelerate the litter decomposition and nutrient cycling compared with Eucalyptus.grandis. Therefore, it will accelerate the. cycle-of matter, improve the soil fertility and stimulate the growth of plant, when Alnus formosana is planted in the plantation of Eucalyptus grandis by the ratio of TⅡ.
(1)桉-桤不同混合比例凋落物中优势类群为弹尾目(Collembola)和蜱螨目(Acariformes)。常见类群为大型土壤动物后孔寡毛目(Opithopora)。五种不同混合比例的凋落物中,大型土壤动物的分布是TV最多,TⅠ最少,混合凋落物介于二者之间,且随桤木比例的增加呈增多的趋势；而中小型土壤动物在TⅠ中最多,而TV最少。啮虫目(Psocoptera)在TV中的数量远远高于其它四种凋落物,后孔寡毛目在混合凋落物中数量较高。弹尾目、蝎线纲(Pauropoda)更喜生活在TⅠ凋落物中。综上表明在巨桉中加入台湾桤木可改变土壤动物的群落结构。
(2)6目凋落袋中土壤动物的个体数是TⅠ>TⅡ>TⅢ>TⅣ>TV,大型土壤动物的个体数是TⅢ>TⅣ>TⅤ>TⅡ>TⅠ,中小型土壤动物个体数是TⅠ>TⅡ>TⅢ>TⅣ>TⅤ,大型土壤动物类群数在TV最多,混合凋落物次之,TⅠ最少。30目凋落袋中土壤动物个体数TⅠ>TⅡ>TV>TⅣ>TⅢ。260目凋落袋中,土壤动物的个体数是Ⅳ>TⅢ>TⅡ>TⅠ>TⅤ。五种不同比例混合凋落物中的土壤动物个体总数在6目中差异极其显著,在30目和260目差异性不显著。中小型土壤动物个体数量是260目>30目>6目,且在30目与6目中差异显著。6目中中小型土壤动物的个体数远高于大型土壤动物。土壤动物类群数是30目>6目>260目,五种不同比例混合凋落物中的土壤动物类群数在260目差异不显著,在30目差异显著,在6目中差异极其显著。表明相对纯桉而言,桉-桤混和可提高凋落物中大型土壤动物的个体数、类群数,而对中小型土壤动物的个体数和类群数并无影响。
(3)6目凋落袋内凋落物的分解速率为TV>TⅣ>TⅢ>TⅡ>TⅠ,TⅠ凋落物的快速分解期在8月,TⅡ和TⅢ凋落物的快速分解期在7月,而TⅣ和TV凋落物的快速分解期在5月。分解前期TⅡ由于桤木凋落物的加入,大大提高了整个凋落物的分解速度,而TⅣ因桉树凋落物的存在,分解速率有所降低。在2009年10月前,260目和30目凋落物袋中,TⅠ凋落物的分解速率快于TV。近自然状态下(6目中),分解前期(0-45d)由于桤木的加入TⅡ凋落物的分解速度有所提高,也即TⅡ(巨桉7g,台湾桤木3g)混合比例可促进凋落物的分解。
(4)背景土壤动物包括湿生中小型土壤动物、干生中小型土壤动物和大型土壤动物。湿生中小型土壤动物中线虫纲(Nematoda)为优势类群,轮虫纲(Rotatoria)和线蚓科(Enchytraeidae)为常见类群。干生中小型土壤动物优势类群有弹尾目和蜱螨目。大型土壤动物优势类群有后孔寡毛目、等翅目(Ⅰsoptera)、鞘翅目(Coleoptera)和膜翅目(Hymenoptera)。在五种类型的人工林中,湿生中小型土壤动物、干生中小型土壤动物以及大型土壤动物的个体数在TI人工林下中均较多,而在混交林和TV林下较少。土壤动物在不同土层个体数差异较大,尤其是大型土壤动物,表聚现象明显。
(5)不同比例桉-桤混合初始凋落物间氮差异极其显著,钙、镁差异显著,碳、磷、钾差异不显著。巨桉、台湾桤木两种凋落物间C：N比差异极其显著。在分解过程中,6目凋落袋中桉-桤不同混合比例凋落物各元素的含量都有所降低。C、K元素含量在TⅠ凋落物中下降最慢,Mg元素含量在TⅠ中下降最快。N、P元素含量在TV凋落物中下降最慢,C、Ca元素含量在TV中下降最快。30目凋落袋中N元素含量在TV凋落物中,C、Ca在TⅠ凋落物中下降缓慢,P、Mg元素含量在TⅠ凋落物中下降较快。260目凋落袋中N、Mg元素含量在TⅠ凋落物中下降迅速,在TV中下降缓慢。6目中凋落物的损失率与N元素、Ca元素的含量呈极其显著正相关,与P元素的含量呈显著正相关,与C含量呈负相关,但相关性不显著。
(6)260目凋落袋中凋落物的养分含量总体上高于30目，也即260目凋落物袋中凋落物的养分释放速率慢于30目,而260目的土壤动物个体数高于30目,土壤动物的类群数低于30目。表明不同的土壤动物类群有利于促进凋落物中养分元素的释放。大型土壤动物对各元素释放作用的大小依次为：Mg>P>K>N>Ca>C,Mg元素释放作用在TV凋落物最高,达92.45%,对C元素释放作用在TⅢ凋落物中最低,仅32.05%,对N元素的释放作用在TⅠ凋落物中最高。
研究表明：不同比例桉-桤混合可提高凋落物中大型土壤动物的个体数与类群数,加速凋落物的分解,促进养分的释放与循环。因此在桉树纯林中混栽桤木(比例为7：3),有利于加速人工林内的物质循环,提高土壤地力,促进植物生长。
Litter decomposition plays an essential role in maintaining soil fertility in the forest. Soil fauna which affected by the type and quality of litter and the stage of decomposition are active participants during the decomposition of litter. Therefore, the research on dynamics of soil fauna community during litter decay can provide important scientific basis for the cultivation and the sustainable management of plantation. Eucalyptus(Eucalyptus grandis) is one of main afforestation tree species in the industrial raw material forest in the south, and alder (Alnus formosana) is a nonleguminous and evergreen tree species with the characteristic of fast growing and nitrogen-fixing. In order to maximum extently avoid the ecological problems, such as soil fertility and biodiversity decline, the litter mixed experiment of Eucalyptus grandis and Alnus formosana was conducted. The corresponding problems were put up:(1) How do control the ratio of eucalyptus litter weight to alder litter weight to maintain productivity and biodiversity in the mixed forest?(2) Will different tree species mixture affect the litter decomposition process?(3) Will different tree species mixture change dynamitic of soil fauna community? All those are lack of corresponding research. Decomposition characteristic and dynamic of soil fauna in different ratio mixed litters (TⅠ:eucalyptus10g, alder Og; TⅡ:eucalyptus7g, alder3g; TⅢ:eucalyptus5g, alder5g; TⅣ:eucalyptus3g, alder7g; TⅤ:eucalyptus Og, alder10g) were investigated with the method of litterbags in Suji town, Leshan city, Sichuan province. The aim was to explore the effect brought by mixture of eucalyptus and alder on decomposition rate, dynamics of nutrient and soil fauna community structure and provide the support of science and technology for the building of reasonable and feasible plantation.
(1) The dominant groups were Collembola and Acariformes which belong to meso and microfauna. The common group was Opithopora which belongs to macrofauna in different ratio mixed litter. Compared with TI litter, the macrofauna preferred to live in TV litter. The individual numbers of macrofauna in mixed litter were between TV litter and TI litter. The individual numbers of macrofauna increases with quantity of the alder added. Compared with TV litter, the meso and microfauna preferred to live in TI litter. The individual numbers of Psocoptera reached highest in TV litter and Opithopora in mixed litter. Collembola and Pauropoda preferred living in TI litter. All these suggested: the community structure of soil fauna would change when alder was added in Eucalyptus.
(2) The individual numbers of soil fauna in6-mesh litterbags were TI>TII>TIII>TIV>TV, and those of macrofauna were TIII>TIV>TV>TII>TI and meso and microfauna were TⅠ>TⅡ>TⅢ>TⅣ>TⅤ. The group numbers of macrofauna were maximum in TV litter and minimum in TI litter. The individual numbers of soil fauna in30-mesh litterbags were TI>TII> TV>TIV>TIII and in260-mesh litterbags were IV> TⅢ>TⅡ>TⅠ>TⅤ. The total individuals of soil fauna showed extremely significant difference in6-mesh litterbags, no significant difference in260-mesh litterbags and30-mesh litterbags among five different ratio mixed litter. The individual numbers of meso and micro fauna were260-mesh>30-mesh>6-mesh and there was significant difference between30-mesh and6-mesh litterbags. The individual numbers of meso and microfauna were more than those of marofauna in6-mesh litterbags. The group numbers were30-mesh>6-mesh>260-mesh and there was significant difference in30-mesh litterbags, extremely significant difference in6-mesh litterbags and no significant difference in260-mesh litterbags among five different ratio mixed litter. All these showed the mixture of eucalyptus and alder could increase the individuals of soil macrofauna, whereas no effect on the individuals and groups of meso and microfauna, compared with the eucalyptus litter.
(3) The decomposition rate was TV>TIV>TIII>TII>TI in6-mesh litterbags, and the fast decomposition stage of TI was in August, TⅡ and TⅢ in July and TIV and TV in May. The decomposition rate of TⅡ litter increased greatly due to the addition of alder in early stage, and the decomposition rate of TIV litter declined because of the eucalyptus added. Before October2009, the decomposition rate of TI litter was faster than that of TⅤ litter in260-and30-mesh litterbags. Under near natural state (6-mesh), the ratio of TⅡ could accelerate the decay of litter.
(4) Background soil fauna included damp living meso and micro fauna, dry living meso and micro fauna and-macrofauna. The-dominant-group was Nematoda and the common groups were Rotatoria and Enchytraeidae among damp living meso and micro fauna. The dominant groups were Collembola and Acariformes among dry living meso-, micro fauna. The dominant groups were Opithopora, Isoptera, Coleoptera and Hymenoptera among macrofauna. There were more individuals in soil under TI plantation than mixed and TV plantation. The soil fauna, especially macrofauna, showed significant difference among different soil layer and more soil fauna happened in the top soil.
(5) There was extremely significant difference in initial N, significant difference in initial Ca and Mg, and no significant difference in C, P and K among different ratio mixed litter. The ratio of C:N showed extremely significant difference between eucalyptus and alder. The elements concentration of different ratio mixed littler declined in6-mesh litterbags during decomposition. The elements of C, K declined slowly and Mg fast in TI litter. The elements of N, P declined slowly and C, Ca fast in TV litter. The element of N, C and Ca declined slowly in TV, TI litter respectively, P, Mg declined fast in TI litter in30-mesh litterbags. N and Mg declined fast in TI litter and slowly in TV litter in260-mesh litterbags. Mass loss rate was extremely significant positively correlated to the concentration of N, Ca, significant correlated to concentration of P, negatively correlate to the content of C, but there was no obvious difference.
(6) The concentration of elements in260-mesh litterbags was higher than in30-mesh litterbags, that is to say, the nutrient release rate of litter in260-mesh litterbags was slower than in30-mesh litterbags, however, the individual numbers of soil fauna were higher in260-mesh litterbags than in-30-mesh litterbags and the group numbers were on the contrary. Different groups of soil fauna could stimulate the release of nutrient elements. The release rate of nutrient elements which was attribute to the soil macrofauna showed the order:Mg>P>K>N>Ca>C. Action of macrofauna to Mg was highest, achieving92.5%, however to C was lowest (32.05%), to N was highest in TI litter.
All the research suggested that the mixture of Eucalyptus grandis and Alnus formosana could increase the numbers of individual and group of macrofauna, accelerate the litter decomposition and nutrient cycling compared with Eucalyptus.grandis. Therefore, it will accelerate the. cycle-of matter, improve the soil fertility and stimulate the growth of plant, when Alnus formosana is planted in the plantation of Eucalyptus grandis by the ratio of TⅡ.
引文
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