Evolution of soil nematode community after establishment of Haloxylon ammodendron plantations in an arid desert-oasis ecotone

doi:10.7522/j.issn.1000-694X.2023.00091

Journal of Desert Research ›› 2024, Vol. 44 ›› Issue (2): 133-142.DOI: 10.7522/j.issn.1000-694X.2023.00091

Evolution of soil nematode community after establishment of Haloxylon ammodendron plantations in an arid desert-oasis ecotone

Fangjiao An¹(), Yongzhong Su²(), Ziru Niu³, Tingna Liu²

^1.School of Civil Engineering，Lanzhou University of Technology，Lanzhou 730050，China
^2.Northwest Institute of Eco-Environment and Resources，Chinese Academy of Sciences，Lanzhou 730000，China
^3.Shaanxi Provincial Land Engineering Construction Group，Xi'an 710075，China

Received:2023-04-14 Revised:2023-10-24 Online:2024-03-20 Published:2024-03-19
Contact: Yongzhong Su

Abstract

Abstract:

Study on the evolution of soil biodiversity during the growth of sand-fixing Haloxylon ammodendron in desert-oasis ecotone is an important part of understanding the plant-soil interaction in arid areas. Diffserent ages （0-， 3-， 6-， 11-， 19-， 28- and 46-years） of H. ammodendron plantations in Linze desert-oasis ecotone of northwestern China were selected and surface soil （0-10 cm） under and outside the canopy were sampled， to study the changes in the composition and diversity of nematode community after sand-fixing H. ammodendron establishment， and to explore the indicator effect of nematode community on plant-soil system restoration. In this study， a total of 15 genera were identified， Eucephalobus and Acrobeloides were the dominant genera. Bacterial nematodes were the dominant trophic group， accounting for 50.3%-94.1% of nematode numbers. The growth of H. ammodendron significantly increased the total number of soil nematodes， bacterial feeding nematodes and nematode diversity. Nematode enrichment index （EI） and structure index （SI） were less than 50. The soil food web was stable first and then degraded gradually with plantation ages. Redundancy analysis （RDA） indicated that changes in soil chemical properties （EC， C/N， and SOC） signi?cantly affected the distribution of nematode communities， and EC had the highest explanation rate for the total variation of nematode genera （32.7%）. Soil nematode as a biological indicator reveals the evolution process of the plant-soil ecosystem.

Key words: sand-fixing Haloxylon ammodendron plantation, sand-fixation year, soil nematode community, diversity, edge of oasis

CLC Number:

S154

Fangjiao An, Yongzhong Su, Ziru Niu, Tingna Liu. Evolution of soil nematode community after establishment of Haloxylon ammodendron plantations in an arid desert-oasis ecotone[J]. Journal of Desert Research, 2024, 44(2): 133-142.

Figures/Tables 10

References 35

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种植年限/a	pH		电导率EC/（μS·cm^-1）		全碳TC/（g·kg^-1）		全氮TN/（g·kg^-1）		全磷TP/（g·kg^-1）
种植年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	9.28±0.16^Ac	9.28±0.16^Aa	130.27±3.93^Ae	130.27±3.93^Ac	4.98±0.11^Ad	4.98±0.11^Ac	0.28±0.04^Ac	0.28±0.04^Ac	0.46±0.02^Ad	0.46±0.02^Ac
3	10.05±0.13^Aab	9.41±0.08^Ba	306.00±10.39^Ade	130.00±5.20^Bc	5.32±0.03^Ad	5.13±0.20^Abc	0.22±0.00^Ad	0.19±0.01^Ad	0.49±0.01^Acd	0.47±0.01^Ac
6	10.08±0.02^Aab	9.59±0.25^Ba	438.00±91.39^Acd	202.27±68.49^Bbc	6.96±0.53^Ac	6.28±0.19^Ab	0.30±0.01^Ac	0.21±0.02^Ad	0.56±0.02^Acd	0.53±0.04^Abc
11	10.03±0.05^Aab	9.61±0.21^Ba	522.00±100.64^Ac	231.00±51.10^Bab	7.40±0.16^Abc	6.32±0.50^Ab	0.38±0.03^Ab	0.28±0.02^Ac	0.57±0.00^Ac	0.54±0.03^Abc
19	10.09±0.01^Aa	9.57±0.37^Ba	713.67±96.26^Abc	253.10±15.53^Bab	8.43±0.88^Aab	7.63±0.92^Aa	0.38±0.01^Ab	0.33±0.00^Abc	0.68±0.07^Aab	0.65±0.08^Aa
28	9.87±0.20^Ab	9.59±0.23^Ba	721.00±71.34^Ab	287.63±104.60^Bab	8.72±1.65^Aa	7.87±1.34^Aa	0.52±0.01^Aa	0.39±0.07^Aa	0.67±0.08^Aabc	0.64±0.08^Aa
46	9.98±0.12^Aab	9.47±0.15^Ba	987.00±227.66^Aa	321.67±45.01^Ba	9.64±0.23^Aa	8.29±0.78^Aa	0.52±0.03^Aa	0.38±0.02^Aab	0.70±0.10^Aa	0.63±0.06^Aab
年限	***	ns	***	*	***	***	***	***	***	*
位置	***		***		ns		ns		ns
年限×位置	***		***		ns		ns		ns
种植年限/a	土壤有机碳SOC/（g·kg^-1）		碳氮比C/N		速效氮AN/（mg·kg^-1）		速效磷AP/（mg·kg^-1）		速效钾AK/（mg·kg^-1）
种植年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	0.35±0.03^Ad	0.35±0.03^Ad	1.29±0.11^Ad	1.29±0.11^Ac	25.38±2.13^Ab	25.38±2.13^Abc	2.33±0.29^Ac	2.33±0.29^Ad	103.33±5.77^Ad	103.33±5.77^Ad
3	0.42±0.04^Ad	0.30±0.02^Bd	1.95±0.19^Ad	1.57±0.04^Ac	20.96±0.52^Bcd	21.93±0.16^Ac	2.52±0.50^Ac	2.79±0.18^Ad	120.00±0.00^Ad	96.67±5.77^Bd
6	1.22±0.09^Ac	0.71±0.13^Bcd	4.02±0.41^Ac	3.39±0.87^Ab	19.63±0.65^Bd	22.81±1.62^Ac	3.40±0.81^Ac	3.56±0.89^Acd	210.00±65.57^Ac	130.00±17.32^Bd
11	1.63±0.13^Ab	1.14±0.19^Bbc	4.38±0.66^Abc	4.06±0.78^Ab	19.86±1.15^Bd	29.56±1.56^Aab	13.37±0.46^Ab	5.84±2.12^Bbc	353.33±41.63^Ab	156.67±15.28^Bcd
19	1.89±0.07^Ab	1.31±0.22^Bb	4.91±0.14^Aab	4.04±0.62^Ab	23.04±0.91^Bbcd	30.26±1.21^Aab	13.94±4.70^Ab	6.45±1.52^Bb	380.00±0.00^Ab	200.00±20.00^Bc
28	2.80±0.54^Aa	1.85±0.62^Ba	5.33±0.92^Aa	4.63±0.72^Aab	24.33±0.73^Bbc	30.84±1.26^Aab	26.85±3.47^Aa	11.92±2.40^Ba	510.00±60.83^Aa	300.00±26.46^Bb
46	2.76±0.13^Aa	2.03±0.20^Ba	5.28±0.43^Aa	5.37±0.82^Aa	29.39±1.09^Ba	33.76±1.52^Aa	23.45±3.29^Aa	11.11±2.09^Ba	546.67±25.17^Aa	453.33±94.52^Ba
年限	***	***	***	***	*	*	***	***	***	***
位置	***		ns		**		***		***
年限×位置	***		ns		ns		*		ns

种植年限/a	pH		电导率EC/（μS·cm^-1）		全碳TC/（g·kg^-1）		全氮TN/（g·kg^-1）		全磷TP/（g·kg^-1）
种植年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	9.28±0.16^Ac	9.28±0.16^Aa	130.27±3.93^Ae	130.27±3.93^Ac	4.98±0.11^Ad	4.98±0.11^Ac	0.28±0.04^Ac	0.28±0.04^Ac	0.46±0.02^Ad	0.46±0.02^Ac
3	10.05±0.13^Aab	9.41±0.08^Ba	306.00±10.39^Ade	130.00±5.20^Bc	5.32±0.03^Ad	5.13±0.20^Abc	0.22±0.00^Ad	0.19±0.01^Ad	0.49±0.01^Acd	0.47±0.01^Ac
6	10.08±0.02^Aab	9.59±0.25^Ba	438.00±91.39^Acd	202.27±68.49^Bbc	6.96±0.53^Ac	6.28±0.19^Ab	0.30±0.01^Ac	0.21±0.02^Ad	0.56±0.02^Acd	0.53±0.04^Abc
11	10.03±0.05^Aab	9.61±0.21^Ba	522.00±100.64^Ac	231.00±51.10^Bab	7.40±0.16^Abc	6.32±0.50^Ab	0.38±0.03^Ab	0.28±0.02^Ac	0.57±0.00^Ac	0.54±0.03^Abc
19	10.09±0.01^Aa	9.57±0.37^Ba	713.67±96.26^Abc	253.10±15.53^Bab	8.43±0.88^Aab	7.63±0.92^Aa	0.38±0.01^Ab	0.33±0.00^Abc	0.68±0.07^Aab	0.65±0.08^Aa
28	9.87±0.20^Ab	9.59±0.23^Ba	721.00±71.34^Ab	287.63±104.60^Bab	8.72±1.65^Aa	7.87±1.34^Aa	0.52±0.01^Aa	0.39±0.07^Aa	0.67±0.08^Aabc	0.64±0.08^Aa
46	9.98±0.12^Aab	9.47±0.15^Ba	987.00±227.66^Aa	321.67±45.01^Ba	9.64±0.23^Aa	8.29±0.78^Aa	0.52±0.03^Aa	0.38±0.02^Aab	0.70±0.10^Aa	0.63±0.06^Aab
年限	***	ns	***	*	***	***	***	***	***	*
位置	***		***		ns		ns		ns
年限×位置	***		***		ns		ns		ns
种植年限/a	土壤有机碳SOC/（g·kg^-1）		碳氮比C/N		速效氮AN/（mg·kg^-1）		速效磷AP/（mg·kg^-1）		速效钾AK/（mg·kg^-1）
种植年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	0.35±0.03^Ad	0.35±0.03^Ad	1.29±0.11^Ad	1.29±0.11^Ac	25.38±2.13^Ab	25.38±2.13^Abc	2.33±0.29^Ac	2.33±0.29^Ad	103.33±5.77^Ad	103.33±5.77^Ad
3	0.42±0.04^Ad	0.30±0.02^Bd	1.95±0.19^Ad	1.57±0.04^Ac	20.96±0.52^Bcd	21.93±0.16^Ac	2.52±0.50^Ac	2.79±0.18^Ad	120.00±0.00^Ad	96.67±5.77^Bd
6	1.22±0.09^Ac	0.71±0.13^Bcd	4.02±0.41^Ac	3.39±0.87^Ab	19.63±0.65^Bd	22.81±1.62^Ac	3.40±0.81^Ac	3.56±0.89^Acd	210.00±65.57^Ac	130.00±17.32^Bd
11	1.63±0.13^Ab	1.14±0.19^Bbc	4.38±0.66^Abc	4.06±0.78^Ab	19.86±1.15^Bd	29.56±1.56^Aab	13.37±0.46^Ab	5.84±2.12^Bbc	353.33±41.63^Ab	156.67±15.28^Bcd
19	1.89±0.07^Ab	1.31±0.22^Bb	4.91±0.14^Aab	4.04±0.62^Ab	23.04±0.91^Bbcd	30.26±1.21^Aab	13.94±4.70^Ab	6.45±1.52^Bb	380.00±0.00^Ab	200.00±20.00^Bc
28	2.80±0.54^Aa	1.85±0.62^Ba	5.33±0.92^Aa	4.63±0.72^Aab	24.33±0.73^Bbc	30.84±1.26^Aab	26.85±3.47^Aa	11.92±2.40^Ba	510.00±60.83^Aa	300.00±26.46^Bb
46	2.76±0.13^Aa	2.03±0.20^Ba	5.28±0.43^Aa	5.37±0.82^Aa	29.39±1.09^Ba	33.76±1.52^Aa	23.45±3.29^Aa	11.11±2.09^Ba	546.67±25.17^Aa	453.33±94.52^Ba
年限	***	***	***	***	*	*	***	***	***	***
位置	***		ns		**		***		***
年限×位置	***		ns		ns		*		ns

固沙年限/a	线虫总数		食细菌线虫BF		食真菌线虫FF		植物寄生线虫PP		捕食/杂食线虫OP
固沙年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	5.33±1.31^Ad	5.33±1.31^Ab	4.00±0.00^Ad	4.00±0.00^Ab	1.33±0.31^Ab	1.33±0.31^Aab	0.00±0.00^Aa	0.00±0.00^Ab	0.00±0.00^Ab	0.00±0.00^Ab
3	44.33±10.18^Acd	11.67±3.15^Bab	36.33±11.16^Acd	8.33±3.15^Bab	6.67±0.58^Aab	1.00±0.73^Bb	0.00±0.00^Aa	2.33±0.08^Aa	1.33±0.31^Ab	0.00±0.00^Ab
6	99.33±22.72^Abc	61.33±11.67^Ba	85.67±24.53^Abc	47.00±9.04^Bab	8.67±1.57^Aab	8.33±1.79^Ba	0.00±0.00^Aa	1.00±0.73^Aab	5.00±1.73^Ab	5.00±1.73^Aab
11	143.33±45.52^Aab	62.00±12.05^Ba	116.00±23.81^Ab	52.33±13.14^Ba	6.33±0.69^Aab	7.33±1.51^Bab	0.00±0.00^Aa	0.00±0.00^Ab	20.00±6.00^Aa	2.33±1.04^Ab
19	148.67±43.94^Aab	61.67±12.53^Ba	123.00±28.58^Ab	49.00±12.26^Bab	8.67±1.77^Aab	1.33±0.31^Bab	1.67±0.89^Aa	1.00±0.73^Aab	15.33±5.15^Aa	10.33±2.45^Aa
28	136.33±40.34^Ab	60.00±11.68^Ba	129.00±43.38^Ab	54.33±18.46^Ba	6.00±1.29^Aab	3.33±0.77^Bab	1.33±0.31^Aa	0.00±0.00^Ab	0.00±0.00^Ab	2.33±1.04^Ab
46	228.00±61.46^Aa	41.00±16.64^Bab	213.33±54.80^Aa	38.67±10.34^Bab	13.67±2.51^Aa	2.33±0.08^Bab	0.00±0.00^Aa	0.00±0.00^Ab	1.00±0.73^Ab	0.00±0.00^Ab
年限	**	*	**	*	ns	ns	ns	ns	*	*
位置	***		***		*		ns		ns
年限× 位置	*		**		ns		ns		ns

固沙年限/a	线虫总数		食细菌线虫BF		食真菌线虫FF		植物寄生线虫PP		捕食/杂食线虫OP
固沙年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	5.33±1.31^Ad	5.33±1.31^Ab	4.00±0.00^Ad	4.00±0.00^Ab	1.33±0.31^Ab	1.33±0.31^Aab	0.00±0.00^Aa	0.00±0.00^Ab	0.00±0.00^Ab	0.00±0.00^Ab
3	44.33±10.18^Acd	11.67±3.15^Bab	36.33±11.16^Acd	8.33±3.15^Bab	6.67±0.58^Aab	1.00±0.73^Bb	0.00±0.00^Aa	2.33±0.08^Aa	1.33±0.31^Ab	0.00±0.00^Ab
6	99.33±22.72^Abc	61.33±11.67^Ba	85.67±24.53^Abc	47.00±9.04^Bab	8.67±1.57^Aab	8.33±1.79^Ba	0.00±0.00^Aa	1.00±0.73^Aab	5.00±1.73^Ab	5.00±1.73^Aab
11	143.33±45.52^Aab	62.00±12.05^Ba	116.00±23.81^Ab	52.33±13.14^Ba	6.33±0.69^Aab	7.33±1.51^Bab	0.00±0.00^Aa	0.00±0.00^Ab	20.00±6.00^Aa	2.33±1.04^Ab
19	148.67±43.94^Aab	61.67±12.53^Ba	123.00±28.58^Ab	49.00±12.26^Bab	8.67±1.77^Aab	1.33±0.31^Bab	1.67±0.89^Aa	1.00±0.73^Aab	15.33±5.15^Aa	10.33±2.45^Aa
28	136.33±40.34^Ab	60.00±11.68^Ba	129.00±43.38^Ab	54.33±18.46^Ba	6.00±1.29^Aab	3.33±0.77^Bab	1.33±0.31^Aa	0.00±0.00^Ab	0.00±0.00^Ab	2.33±1.04^Ab
46	228.00±61.46^Aa	41.00±16.64^Bab	213.33±54.80^Aa	38.67±10.34^Bab	13.67±2.51^Aa	2.33±0.08^Bab	0.00±0.00^Aa	0.00±0.00^Ab	1.00±0.73^Ab	0.00±0.00^Ab
年限	**	*	**	*	ns	ns	ns	ns	*	*
位置	***		***		*		ns		ns
年限× 位置	*		**		ns		ns		ns

线虫属	营养类型	c-p值	线虫属的优势度
			0 a		3 a		6 a		11 a		19 a		28 a		46 a
			冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
Mesorhabditis	BF	1	—	—	++	—	++	++	++	++	++	++	—	++	++	—
Eucephalobus	BF	2	+++	+++	+++	++	+++	+++	+++	++	+++	+++	+++	+++	+++	+++
Acrobeles	BF	2	—	—	++	+++	+++	+++	++	+++	++	++	+++	++	++	++
Acrobeloides	BF	2	+++	+++	+++	+++	+++	+++	+++	+++	+++	+++	+++	+++	+++	+++
Eumonhystera	BF	2	—	—	—	—	—	++	—	—	—	—	—	—	—	—
Ceratoplectus	BF	2	—	—	—	—	—	—	—	—	—	—	—	—	++	—
Paraphelenchus	FF	2	+++	+++	++	+++	++	++	++	+++	++	—	++	++	++	++
Aphelenchoides	FF	2	—	—	—	—	++	—	—	—	++	—	—	—	—	—
Tylolaimophorus	FF	3	—	—	++	—	++	+++	++	++	++	++	++	—	+	++
Filenchus	PP	2	—	—	—	+++	—	—	—	—	—	—	—	—	—	—
Psilenchus	PP	2	—	—	—	+++	—	++	—	—	++	—	+	—	—	—
Rotylenchus	PP	3	—	—	—	—	—	—	—	—	++	++	—	—	—	—
Prodorylaimium	OP	5	—	—	—	—	—	—	+	—	++	++	—	—	—	—
Aporcelaimium	OP	5	—	—	++	—	++	++	+++	++	++	+++	—	++	+	—
Discolaimus	OP	5	—	—	—	—	++	—	—	—	—	—	—	—	—	—

Evolution of soil nematode community after establishment of Haloxylon ammodendron plantations in an arid desert-oasis ecotone

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RDA轴	1	2	3	4
蒙特卡罗排列检验	R²=0.476 F=26.2 P=0.002^***
特征值	0.4279	0.0327	0.0106	0.005
累积解释变异/%	42.79	46.06	47.12	47.61
线虫属相关性	0.9333	0.4264	0.3507	0.3215
累计解释拟合变异/%	89.3	96.12	98.33	99.36

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固沙年限/a	丰富度指数		香农-维纳多样性指数		均匀度指数		优势度指数		总成熟度指数		线虫通路指数
固沙年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	0.40±0.06^e	0.40±0.06^c	0.23±0.40^Ab	0.23±0.40^Ac	0.33±0.07^Ab	0.33±0.07^Ab	0.83±0.29^Aa	0.83±0.29^Aa	2.00±0.00^Aa	2.00±0.00^Ab	0.83±0.29^Aa	0.83±0.29^Aa
3	1.33±0.18^bc	1.19±0.32^ab	1.11±0.33^Aa	0.75±0.10^Ab	0.84±0.10^Ba	0.93±0.12^Aa	0.40±0.11^Ab	0.50±0.00^Ab	2.13±0.13^Aa	2.00±0.00^Ab	0.82±0.07^Aa	0.67±0.58^Aa
6	2.04±0.19^a	1.65±0.42^a	1.38±0.16^Aa	1.59±0.08^Aa	0.76±0.01^Ba	0.88±0.08^Aa	0.33±0.04^Ab	0.40±0.07^Abc	2.15±0.09^Aa	2.39±0.24^Aa	0.89±0.10^Aa	0.74±0.30^Aa
11	1.38±0.23^bc	1.30±0.36^ab	1.17±0.18^Aa	0.96±0.38^Ab	0.70±0.08^Bab	0.81±0.08^Aa	0.48±0.15^Ab	0.38±0.14^Abc	2.33±0.60^Aa	2.04±0.14^Ab	0.96±0.04^Aa	0.86±0.17^Aa
19	1.42±0.02^b	1.31±0.25^ab	1.30±0.09^Aa	1.22±0.34^Aab	0.72±0.03^Bab	0.77±0.05^Aa	0.32±0.06^Ab	0.24±0.02^Ac	2.42±0.09^Aa	2.45±0.13^Aa	0.94±0.05^Aa	0.98±0.03^Aa
28	0.96±0.26^cd	0.95±0.33^abc	1.12±0.11^Aa	0.95±0.39^Ab	0.77±0.09^Ba	0.78±0.15^Aa	0.40±0.07^Ab	0.48±0.18^Abc	2.01±0.02^Aa	2.06±0.10^Ab	0.94±0.06^Aa	0.92±0.14^Aa
46	0.87±0.10^d	0.86±0.24^bc	1.07±0.07^Aa	0.81±0.23^Ab	0.64±0.03^Bab	0.78±0.15^Aa	0.45±0.03^Ab	0.53±0.12^Ab	2.00±0.05^Aa	2.02±0.04^Ab	0.94±0.01^Aa	0.92±0.09^Aa
年限	***	ns	*	*	*	*	*	*	ns	**	ns	ns
位置	ns		ns		**		ns		ns		ns
年限× 位置	ns		ns		ns		ns		ns		ns

固沙年限/a	丰富度指数		香农-维纳多样性指数		均匀度指数		优势度指数		总成熟度指数		线虫通路指数
固沙年限/a	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外	冠层下	冠层外
0	0.40±0.06^e	0.40±0.06^c	0.23±0.40^Ab	0.23±0.40^Ac	0.33±0.07^Ab	0.33±0.07^Ab	0.83±0.29^Aa	0.83±0.29^Aa	2.00±0.00^Aa	2.00±0.00^Ab	0.83±0.29^Aa	0.83±0.29^Aa
3	1.33±0.18^bc	1.19±0.32^ab	1.11±0.33^Aa	0.75±0.10^Ab	0.84±0.10^Ba	0.93±0.12^Aa	0.40±0.11^Ab	0.50±0.00^Ab	2.13±0.13^Aa	2.00±0.00^Ab	0.82±0.07^Aa	0.67±0.58^Aa
6	2.04±0.19^a	1.65±0.42^a	1.38±0.16^Aa	1.59±0.08^Aa	0.76±0.01^Ba	0.88±0.08^Aa	0.33±0.04^Ab	0.40±0.07^Abc	2.15±0.09^Aa	2.39±0.24^Aa	0.89±0.10^Aa	0.74±0.30^Aa
11	1.38±0.23^bc	1.30±0.36^ab	1.17±0.18^Aa	0.96±0.38^Ab	0.70±0.08^Bab	0.81±0.08^Aa	0.48±0.15^Ab	0.38±0.14^Abc	2.33±0.60^Aa	2.04±0.14^Ab	0.96±0.04^Aa	0.86±0.17^Aa
19	1.42±0.02^b	1.31±0.25^ab	1.30±0.09^Aa	1.22±0.34^Aab	0.72±0.03^Bab	0.77±0.05^Aa	0.32±0.06^Ab	0.24±0.02^Ac	2.42±0.09^Aa	2.45±0.13^Aa	0.94±0.05^Aa	0.98±0.03^Aa
28	0.96±0.26^cd	0.95±0.33^abc	1.12±0.11^Aa	0.95±0.39^Ab	0.77±0.09^Ba	0.78±0.15^Aa	0.40±0.07^Ab	0.48±0.18^Abc	2.01±0.02^Aa	2.06±0.10^Ab	0.94±0.06^Aa	0.92±0.14^Aa
46	0.87±0.10^d	0.86±0.24^bc	1.07±0.07^Aa	0.81±0.23^Ab	0.64±0.03^Bab	0.78±0.15^Aa	0.45±0.03^Ab	0.53±0.12^Ab	2.00±0.05^Aa	2.02±0.04^Ab	0.94±0.01^Aa	0.92±0.09^Aa
年限	***	ns	*	*	*	*	*	*	ns	**	ns	ns
位置	ns		ns		**		ns		ns		ns
年限× 位置	ns		ns		ns		ns		ns		ns