黄土丘陵区不同立地条件下猪毛蒿种内、种间竞争

doi:10.17521/cjpe.2006.0079

摘要/Abstract

摘要：

猪毛蒿(Artemisia scoparia)通常是黄土丘陵区撂荒演替前期群落优势种,在无人为干扰的情况下,猪毛蒿群落通常会向冰草(Agropyron cristatum)群落或阿尔泰狗娃花(Heteropappus altaicus)群落,或长芒草(Stipa bungeana)群落等演替。该文通过河阶地和梁峁阴坡中猪毛蒿生长特征的调查和种内、种间竞争田间试验,从植物竞争角度对猪毛蒿群落的演替机制给予了解释。结果表明:1)两类样地中猪毛蒿的生长都趋于小型化,有少数大个体和多数小个体,都存在异速生长现象,说明两类样地存在竞争,且对猪毛蒿的生长形态具有塑造作用; 2)由于单位地上生物量竞争效应排除了立地条件和个体大小的影响,因而比总竞争效应更能说明种对间的相对竞争能力。梁峁阴坡地和一、二级河阶地三种立地条件下,7种测试植物中对猪毛蒿的相对竞争能力以演替后期多年生植物较高,而演替前期一年生植物较低,说明演替后期种对前期种的竞争抑制是演替驱动力之一; 3)以各测试植物对猪毛蒿单位重量竞争抑制程度平均值来看,以梁峁阴坡地最大,二级河阶地次之,一级河阶地最小,说明立地条件越差,土壤资源可利用水平越低,竞争越激烈; 4)一级河阶地和梁峁阴坡地各测试植物对猪毛蒿的竞争等级发生了显著变化,说明环境条件差别较大时,植物的竞争等级会发生变化。

关键词: 猪毛蒿, 种间竞争, 种内竞争, 异速生长

Abstract:

Background and Aims In hilly Loess region, Artemisia scoparia is always the dominator in the early abandoned field communities. It may be replaced by Agropyron cristatum or Heteropappus altaicus or Stipa bungeana in next succession stage when no disturbance occurs. In order to study the direction of Artemisia scoparia community succession under different standing conditions, we conducted transplant experiments to test the intraspecfic and interspecfic competition of Artemisia scoparia in river terrace and northern mound land and, at the same time, its morphological traits were investigated.
Methods In the experiments, nine species (three coexisting species, five later sere species and Artemisia scoparia itself) were chosen and transplanted as test species, but due to the low survival of two species, only seven species were analyzed to test the relative competition ability to phytometer Artemisia scoparia under different standing conditions.
Key Results The results show that the individuals tend to be miniaturized, i.e., its communities are composed mainly of small-sized individuals and competition can change the morphological traits. The unit biomass competition intensity is better than total competition intensity in explaining plants' relative competitive ability when local conditions and plant size are considered. Under the northern mound land and the lowest river terrace, late-successional and perennials species have higher competitative ability than early-successional and annual plants, suggesting that the competition of late- against early-successional species is one of the driving force to succession. Comparison of the competition between the seven test species and the unit biomass of Artemisia scoparia, the northern mound land ranks the first, lower river terrace the second and the lowest terrace the third, suggesting that intensive competition occurs more severely in poor growing conditions. There is significant difference in the ability of competition between the test species and Artemisia scoparia from the lowest river terrace to the northern mound land, suggesting that the ranks of the competition ability may vary depending on standing conditions.
Conclusions As competition is one of the decisive factors in constructing plant community, the shift or variation of competition ability between coexisting species and sere species may result in different community structure and dynamics, so it changes the direction and pathway of succession under different standing conditions.

Key words: Artemisia scoparia, Interspecific competition, Intraspecfic competition, Allometry

杜峰, 梁宗锁, 山仑, 陈小燕. 黄土丘陵区不同立地条件下猪毛蒿种内、种间竞争. 植物生态学报, 2006, 30(4): 601-609. DOI: 10.17521/cjpe.2006.0079

DU Feng, LIANG Zong-Suo, SHAN Lun, CHEN Xiao-Yan. INTRASPECFIC AND INTERSPECFIC COMPETITION OF ARTEMISIA SCOPARIA UNDER DIFFERENT SITE CONDITIONS IN THE HILLY REGION OF LOESS PLATEAU. Chinese Journal of Plant Ecology, 2006, 30(4): 601-609. DOI: 10.17521/cjpe.2006.0079

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2006.0079

https://www.plant-ecology.com/CN/Y2006/V30/I4/601

图/表 5

表1 研究地点土壤水分、养分概况

Table 1 Summary of soil moisture and nutrition in investigation and experiment sites

调查和试验地点 Investigation and experiment sites	土壤水分含量(%) Soil moisture content					土壤养分 Soil nutrients
调查和试验地点 Investigation and experiment sites	0~20 cm	20~40 cm		40~60 cm		有机质 Organic (g·kg^-1)	全氮 Total N (g·kg^-1)	速效氮 Active N (mg·kg^-1)	速效磷 Active P (mg·kg^-1)	速效钾 Active K (mg·kg^-1)
新庄科一级河阶地 The lowest river terrace in Xinzhuangke	10.55		10.12		11.15	7.86	0.77	20.36	4.89	122.16
千腰岘梁峁阴坡 Northern mound land in Qianyaoxian	6.04		7.28		7.74	5.42	0.39	16.45	2.08	74.62
仁台沟一级河阶地 The lowest river terrace in Rentaihou	14.76		15.20		15.83	11.25	0.63	20.21	1.66	71.01
仁台沟二级河阶地 Lower river terrace in Rentaigou	9.06		10.79		11.28	7.52	0.59	17.08	1.87	68.43

图1 不同立地条件下猪毛蒿高度和地上生物量频次分布图图中左侧为河阶地,右侧为梁峁坡地

Fig.1 Frequency histogram of height and aboveground biomass of Artemisia scoparia under different standing conditions In the figure, left side is river terrace old-field and right side is northern mound old-field

图2 猪毛蒿非线形异速生长LogH-LogB平面图

Fig.2 LogH-LogB graph of nonlinear allometry of Artemisia scoparia

表2 不同立地条件下猪毛蒿种内、种间竞争及其测试植物的生长特征

Table 2 Growth traits of Artemisia scoparia with and without test species

地点	测试植物 Test species			目标种 Phytometer				样本数N
Sites	植物种名	地上生物量(FWg/穴)		高度		地上生物量(FWg/穴)
	Species	Aboveground biomass		Height (cm)		Aboveground biomass
		per bunch		Height (cm)		per bunch
		平均	标准差	平均	标准差	平均	标准差
		Mean	SD	Mean	SD	Mean	SD
梁峁阴坡	无 No			100.49	28.66	255.58	110.46	27
Northern
mound land
	狗尾草Setaria viridis	6.2	1.82	94.14	27.85	238.45	99.47	15
	猪毛蒿 Artemissia scoparia	210.84	135.41	45.82	24.03	14.42	9.79	22
	冰草Agropyron cristatum	2.23	2	98.18	20.01	212.97	86.56	25
	阿尔泰狗娃Heteropappus altaicus	32.54	25.53	72.11	14.67	117.3	87.66	19
	长芒草Stipa bungeana	5.46	3.85	87.25	15.89	189.86	91.47	13
	中华隐子草 Ceistogenes squarrosa	42.51	26.13	42.2	15.4	60.07	30.2	16
	白羊草Bothriochloa ischaemum	67.89	52.57	38.05	1756	41.24	27.38	12
一级河阶地	无 No			91.56	31.99	243.19	139.01	13
The lowest river
treeace
	狗尾草S. viridis	3.58	2.56	88.87	27.58	240.07	120.33	12
	猪毛蒿 A. scoparia	134.25	78.89	71.26	31.07	48.36	22.52	9
	冰草A. cristatum	6.46	4.19	88.47	35.78	200.54	130.13	8
	阿尔泰狗娃H. altaicus	30.45	20.05	78.65	33.6	112.23	64.7	12
	长芒草S. bungeana	20.47	13.88	79.46	25.66	160.27	90.52	11
	中华隐子草 C.squarrosa	30.51	10.42	50.26	25.09	76.12	40.56	12
	白羊草B. ischaemum	30.55	22.37	53.63	28.32	69.46	35.04	8
二级河阶地	无 No			129.11	23.79	277.67	131.15	10
Lower river
terrace
	狗尾草S. viridis	10.14	3.22	120.15	26.75	246.5	128.62	9
	猪毛蒿 A. scoparia	156.3	75.28	88.49	36.76	38.99	20.18	12
	冰草A. cristatum	8.46	5.18	110.35	26.03	213.85	140.49	9
	阿尔泰狗娃H. altaicus	40.35	24.57	80.17	42.03	97.46	68.74	11
	长芒草S. bungeana	20.2	10.69	96.44	26.35	195.86	103.33	12
	中华隐子草 C. squarrosa	36.9	20.48	39.45	27.26	57.81	39.68	8

表2 不同立地条件下猪毛蒿种内、种间竞争及其测试植物的生长特征

Table 2 Growth traits of Artemisia scoparia with and without test species

地点	测试植物 Test species			目标种 Phytometer				样本数N
Sites	植物种名	地上生物量(FWg/穴)		高度		地上生物量(FWg/穴)
	Species	Aboveground biomass		Height (cm)		Aboveground biomass
		per bunch		Height (cm)		per bunch
		平均	标准差	平均	标准差	平均	标准差
		Mean	SD	Mean	SD	Mean	SD
梁峁阴坡	无 No			100.49	28.66	255.58	110.46	27
Northern
mound land
	狗尾草Setaria viridis	6.2	1.82	94.14	27.85	238.45	99.47	15
	猪毛蒿 Artemissia scoparia	210.84	135.41	45.82	24.03	14.42	9.79	22
	冰草Agropyron cristatum	2.23	2	98.18	20.01	212.97	86.56	25
	阿尔泰狗娃Heteropappus altaicus	32.54	25.53	72.11	14.67	117.3	87.66	19
	长芒草Stipa bungeana	5.46	3.85	87.25	15.89	189.86	91.47	13
	中华隐子草 Ceistogenes squarrosa	42.51	26.13	42.2	15.4	60.07	30.2	16
	白羊草Bothriochloa ischaemum	67.89	52.57	38.05	1756	41.24	27.38	12
一级河阶地	无 No			91.56	31.99	243.19	139.01	13
The lowest river
treeace
	狗尾草S. viridis	3.58	2.56	88.87	27.58	240.07	120.33	12
	猪毛蒿 A. scoparia	134.25	78.89	71.26	31.07	48.36	22.52	9
	冰草A. cristatum	6.46	4.19	88.47	35.78	200.54	130.13	8
	阿尔泰狗娃H. altaicus	30.45	20.05	78.65	33.6	112.23	64.7	12
	长芒草S. bungeana	20.47	13.88	79.46	25.66	160.27	90.52	11
	中华隐子草 C.squarrosa	30.51	10.42	50.26	25.09	76.12	40.56	12
	白羊草B. ischaemum	30.55	22.37	53.63	28.32	69.46	35.04	8
二级河阶地	无 No			129.11	23.79	277.67	131.15	10
Lower river
terrace
	狗尾草S. viridis	10.14	3.22	120.15	26.75	246.5	128.62	9
	猪毛蒿 A. scoparia	156.3	75.28	88.49	36.76	38.99	20.18	12
	冰草A. cristatum	8.46	5.18	110.35	26.03	213.85	140.49	9
	阿尔泰狗娃H. altaicus	40.35	24.57	80.17	42.03	97.46	68.74	11
	长芒草S. bungeana	20.2	10.69	96.44	26.35	195.86	103.33	12
	中华隐子草 C. squarrosa	36.9	20.48	39.45	27.26	57.81	39.68	8

表3 不同立地条件下各测试植物对猪毛蒿的相对竞争强度及其对猪毛蒿生长的影响

Table 3 The significance level of Artemisia scoparia traits under different test species' competition

地点 Sites	测试植物 Test species	高度 Height			地上生物量 Biomass above ground
地点 Sites	测试植物 Test species	P_U	RC $I Total H$		P_U	RC $I Total H$	RCI_Pergram
梁峁阴坡地 Northern mound land	狗尾草Setaria viridis	0.491 2		0.063 2	0.616 6	0.067 0	0.010 8
	猪毛蒿 Artemisia scoparia	0.000 1		0.544 0	0.000 1	0.943 6	0.004 5
	冰草Agropyron cristatum	0.739 5		0.023 0	0.106 3	0.166 7	0.074 8
	阿尔泰狗娃Heteroppus altaicus	0.000 1		0.282 4	0.000 1	0.541 0	0.016 6
	长芒草Stipa bungeana	0.068 7		0.131 8	0.062 9	0.257 1	0.047 1
	中华隐子草 Cleistogenes squarrosa	0.000 1		0.580 1	0.000 1	0.765 0	0.018 0
	白羊草Bothriochloa ischaemum	0.000 1		0.621 4	0.000 1	0.838 6	0.012 4
一级河阶地 The lowest river terrace	狗尾草S. viridis	0.824 5		0.029 4	0.960 2	0.012 8	0.003 6
	猪毛蒿 A. scoparia	0.154 4		0.221 7	0.000 8	0.801 1	0.006 0
	冰草A. cristatum	0.839 2		0.033 7	0.531 8	0.175 4	0.027 1
	阿尔泰狗娃H. altaicus	0.335 3		0.141 0	0.014 7	0.538 5	0.017 7
	长芒草S. bungeana	0.324 1		0.132 2	0.193 5	0.341 0	0.016 7
	中华隐子草 C.squarrosa	0.001 6		0.451 1	0.002 6	0.687 0	0.022 5
	白羊草B. ischaemum	0.012 7		0.414 3	0.002 0	0.714 4	0.023 4
二级河阶地 Lower river terrace	狗尾草S. viridis	0.450 1		0.069 4	0.715 8	0.112 3	0.011 1
	猪毛蒿 A.scoparia	0.031 5		0.314 6	0.003 1	0.859 6	0.005 5
	冰草A. cristatum	0.453 8		0.145 3	0.482 9	0.229 8	0.027 2
	阿尔泰狗娃H. altaicus	0.004 3		0.379 1	0.017 5	0.649 0	0.016 1
	长芒草S. bungeana	0.104 7		0.253 0	0.256 8	0.294 6	0.014 6
	中华隐子草 C.squarrosa	0.000 1		0.694 4	0.004 8	0.791 8	0.021 5

表3 不同立地条件下各测试植物对猪毛蒿的相对竞争强度及其对猪毛蒿生长的影响

Table 3 The significance level of Artemisia scoparia traits under different test species' competition

地点 Sites	测试植物 Test species	高度 Height			地上生物量 Biomass above ground
地点 Sites	测试植物 Test species	P_U	RC $I Total H$		P_U	RC $I Total H$	RCI_Pergram
梁峁阴坡地 Northern mound land	狗尾草Setaria viridis	0.491 2		0.063 2	0.616 6	0.067 0	0.010 8
	猪毛蒿 Artemisia scoparia	0.000 1		0.544 0	0.000 1	0.943 6	0.004 5
	冰草Agropyron cristatum	0.739 5		0.023 0	0.106 3	0.166 7	0.074 8
	阿尔泰狗娃Heteroppus altaicus	0.000 1		0.282 4	0.000 1	0.541 0	0.016 6
	长芒草Stipa bungeana	0.068 7		0.131 8	0.062 9	0.257 1	0.047 1
	中华隐子草 Cleistogenes squarrosa	0.000 1		0.580 1	0.000 1	0.765 0	0.018 0
	白羊草Bothriochloa ischaemum	0.000 1		0.621 4	0.000 1	0.838 6	0.012 4
一级河阶地 The lowest river terrace	狗尾草S. viridis	0.824 5		0.029 4	0.960 2	0.012 8	0.003 6
	猪毛蒿 A. scoparia	0.154 4		0.221 7	0.000 8	0.801 1	0.006 0
	冰草A. cristatum	0.839 2		0.033 7	0.531 8	0.175 4	0.027 1
	阿尔泰狗娃H. altaicus	0.335 3		0.141 0	0.014 7	0.538 5	0.017 7
	长芒草S. bungeana	0.324 1		0.132 2	0.193 5	0.341 0	0.016 7
	中华隐子草 C.squarrosa	0.001 6		0.451 1	0.002 6	0.687 0	0.022 5
	白羊草B. ischaemum	0.012 7		0.414 3	0.002 0	0.714 4	0.023 4
二级河阶地 Lower river terrace	狗尾草S. viridis	0.450 1		0.069 4	0.715 8	0.112 3	0.011 1
	猪毛蒿 A.scoparia	0.031 5		0.314 6	0.003 1	0.859 6	0.005 5
	冰草A. cristatum	0.453 8		0.145 3	0.482 9	0.229 8	0.027 2
	阿尔泰狗娃H. altaicus	0.004 3		0.379 1	0.017 5	0.649 0	0.016 1
	长芒草S. bungeana	0.104 7		0.253 0	0.256 8	0.294 6	0.014 6
	中华隐子草 C.squarrosa	0.000 1		0.694 4	0.004 8	0.791 8	0.021 5

参考文献 40

[1]	Aerts R (1999). Interspecfic competition in natural plant communities: mechanisms, trade-offs and plant-soil feedbacks. Journal of Experimental Botany, 50, 29-37.
[2]	Aguiar MR, Lauenroth WK, Peters DP (2001). Intensity of intra- and interspecific competition in coexisting shortgrass species. Journal of Ecology, 89, 40-47.
[3]	Begon M, Harper JL, Townsend CR (1990). Ecology, Individuals, Populations and Communities 2nd edn. Blackwell Scientific Publications, Oxford.
[4]	Cahill JF, Casper BB (2000). Investigating the relationship between neighbor root biomass and belowground competition: field evidence for symmetric competition belowground. Oikos, 90, 311-320.
[5]	Casper BB, Jackson RB (1997). Plant competition underground. Annual Review of Ecology and Systematics, 28, 545-570.
[6]	Cheng ZH (2001). Growth and Competition of Fourteen Herb species in Experimental Microcoenosiums. Acta Botanica Sinica(植物学报), 43, 1184-1190.
[7]	Crawley MJ, May RM(1987). Population dynamics and plant community structure: competition between annuals and perennials. Journal of Theoretical Biology, 125, 475-489.
[8]	Davies W (1928). The factor of competition between one species and another in seeds mixtures. In: Welsh Plant Breeding Station ed. Bulletin H8. Aberystwyth, 82-149.
[9]	Du F( 杜峰), Liang ZS( 梁宗锁), Hu LJ ( 胡莉娟) (2004). A reviews on plant competition. Chinese Journal of Ecology(生态学杂志), 23(4), 157-163. (in Chinese with English abstract)
[10]	Firbank LG, Watkinson AR(1990). On the effects of competition: from monocultures to mixtures. In: Grace JB, Tilman D eds. Perspectives on Plant Competition. Academic Press, New York, 165-192.
[11]	Fowler N (1986). The role of competition in plant communities in arid and semiarid regions. Annual Review of Ecology and Systematics, 17, 89-110.
[12]	Goldberg DE (1990). Components of resources in plant communities. In: Grace JB, Tilman D eds. Perspectives on the Plant Competition. Academic Press, New York, 27-49.
[13]	Goldberg DE (1996). Competitive ability: definitions contingency and correlated traits. Philosophical Transactions of the Royal Society of London B, 351, 1377-1385.
[14]	Grace JB (1989). Effects of water depth on Typha latifolia and Typha domingensis. American Journal of Botany, 76, 762-768.
[15]	Grace JB (1995). On the measurement of plant competition intensity. Ecology, 76, 305-308.
[16]	Hamilton JG, Holzapfel C, Mahall BE (1999). Coexistence and interference between a native perennial grass and non-native annaul grasses in California. Oecologia, 121, 518-526. URL PMID
[17]	Howard TG (2001). The relationship of total and per-gram rankings in competitive effect to the natural abundance of herbaceous perennials. Journal of Ecology, 89, 110-117.
[18]	Keddy P, Nielsen K, Weiher E, Lawson R (2002). Relative competitive performance of 63 species of terrestrial herbaceous plants. Journal of Vegetation Science, 13, 5-16.
[19]	Leuschner C, Hertel D, Coners H, Büttner V (2001). Root competition between beech an oak:a hypothesis. Oecologia, 126, 276-284. URL PMID
[20]	Li B( 李博), Chen JK( 陈家宽), Watkinson AR (1998). A literature review on plant competition. Chinese Buttetin of Botany(植物学通报), 15(4), 18-29. (in Chinese with English abstract)
[21]	Li B( 李博) (2001). Plant Competition—Studies on the Interactions Between Crop and Weed. Higher Education Press and Springer Press, Beijing, 251. (in Chinese)
[22]	Li ZF( 李仲芳), Wang G( 王刚) (2002). The effects of intraspecific competition on the relationship between height growth and biomass of annual plants. Journal of Lanzhou University (Natural Sciences)(兰州大学学报(自然科学版)), 38(2), 141-146. (in Chinese with English abstract)
[23]	Martin RA, William KL, Debrap PP (2001). Intensity of intra- and interspecific competition in coexisting shortgrass species. Journal of Ecology, 89, 40-47.
[24]	Miller TE, Werner PA(1987). Competitive effects and response between plant species in a first year old-field community. Ecology, 68, 1201-1210.
[25]	Negbi M, Evenari M(1961). The means of survival of some desert summer annuals. In: Plant-Water Relationships in Arid and Semiarid Conditions. UNESCO, Paris, 249.
[26]	Rees M, Bergelson J(1997). Asymmetric light competition and founder control in plant communities. Journal of Theoretical Biology, 184, 353-358. DOI URL PMID
[27]	Rees M, Condit R, Crawley M, Pacala S, Tilman D(2001). Long-term studies of vegetation dynamics. Science, 293, 650-655. URL PMID
[28]	Sharitz RR, McCormick JF(1973). Population dynamics of two competition plant annual species. Ecology, 54, 723-740.
[29]	Spillards DM(1989). Studies of Plant Competition. PhD dissertation, University of East Anglia, Norwic.
[30]	Tang QY( 唐启义), Feng MG ( 冯明光) (1997). Practical Statistics and DPS Data Processing System. China Agricultural Press, Beijing, 407. (in Chinese)
[31]	Tielbørger K, Kadmon R(2000). Indirect effects in a desert plant community: is competition among annuals more intense under shrub canopies? Plant Ecology, 150, 53-63.
[32]	Tilman D(1982). Resource Competition and Community Structure. Princeton University Press, Princeton, New Jersey.
[33]	Tilman D(1988). Plant Strategies and the Dynamics and the Structure of Plant Communities. Princeton University Press, Princeton, New Jersey.
[34]	Turner MD, Rabinowitz D(1983). Factors affecting frequency distributions of plant mass: the absence of dominance and suppression in competing monocultures of Festuca paradoxa. Ecology, 64, 469-475.
[35]	Wang ZQ( 王政权), Wu GS( 吴巩胜), Wang JB( 王军邦)(2000). Application of competition index in assessing intraspecfic and interspecfic spatial relations between manchurian ash and dahurian larch. Chinese Journal of Applied Ecology(应用生态学报), 11, 641-645.(in Chinese with English abstract) URL PMID
[36]	Weiner J, Thomas SC(1986). Size variability and competition in plant monocultures. Oikos, 47, 211-222.
[37]	Weiner J, Thomas SC (1992). Competition and allometry in three species of annual plants. Ecology, 73, 648-656.
[38]	Wichmann L(2001). Annual variations in competition symmetry in even-aged sitka spruce. Annals of Botany, 88, 145-151.
[39]	Zhang ZP( 张泽浦), Fang JY( 方精云), Kan M(2000). Effects of competition on growth rate and probability of death of plant individuals: a study based on nursery experiments of Larix leptolepis populations. Acta Phytoecologica Sinica(植物生态学报), 24, 340-345. (in Chinese with English abstract)
[40]	Zou CJ( 邹春静), Xu WD( 徐文铎)(1998). Study on intraspecific and interspecfic competition of Picea mongolica. Acta Phytoecologfca Sinica (植物生态学报), 22, 269-274. (in Chinese with English abstract)