C:N:P stoichiometry across evergreen broad-leaved forests, evergreen coniferous forests and deciduous broad-leaved forests in the Tiantong region, Zhejiang Province, eastern China

doi:10.3773/j.issn.1005-264x.2010.01.008

Abstract

Abstract:

Aims Little is known about constrained ratios of carbon, nitrogen, and phosphorus (C:N:P) in terrestrial ecosystems. Our objective was to examine the C:N:P stoichiometry and its relationship with N and P resorption in evergreen broad-leaved forests (EBLF), evergreen coniferous forests (CF) and deciduous broad-leaved forests (DF) at the regional scale.

Methods The study was conducted in Tiantong National Forest Park (29°52′ N, 121°39′ E), Zhejiang Province, eastern China. To estimate foliar and litter C:N:P ratios and N and P resorption efficiencies, we quantified the C, N and P concentrations in leaf and litterfall in EBLF, CF and DF. We used type II regression slopes (reduced major axis, RMA) to determine whether C:N:P stoichiometry varied across gradients of forest production and nutrients.

Important findings The C:N:P ratios in EBLF, CF and DF were 758:18:1, 678:14:1 and 338:11:1 in fresh leaves and 777:13:1, 691:14:1 and 567:14:1 in litterfall, respectively. The foliar C:N ratio was highest in CF, intermediate in EBLF and lowest in DF, while the foliar C:P and N:P ratios were highest in EBLF, intermediate in CF and lowest in DF. In contrast, the litterfall C:N and C:P ratios were higher in EBLF than in CF and DF, and there were no significant differences of N:P ratio among forests. The type II regression slope for N vs. P in leaves of overall plants was statistically >1, suggesting an increasing investment of N with increasing of P in fresh leaves. In contrast, the slope for N vs. P in litterfall approximated 1. N resorption in EBLF was significantly higher than in CF and in DF, but the highest P resorption was observed in DF. Although foliar N:P ratios indicated that EBLF was P limited, DF was N limited and CF was both N and P limited, the nutrient resorption efficiency did not respond with relatively high N resorption in EBLF and high P resorption in DF. We concluded that the relative higher resorption of N and P before leaf abscission could be an inherent property of plants, but was not a mechanism thought to have evolved to conserve nutrients in environments with limited N or P supply.

Key words: C:N:P ratio, model Type II regression, nutrient limitation, nutrient resorption, stoichiometry

YAN En-Rong, WANG Xi-Hua, GUO Ming, ZHONG Qiang, ZHOU Wu. C:N:P stoichiometry across evergreen broad-leaved forests, evergreen coniferous forests and deciduous broad-leaved forests in the Tiantong region, Zhejiang Province, eastern China[J]. Chin J Plant Ecol, 2010, 34(1): 48-57.

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Figures/Tables 4

References 32

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植被类型和群落类型 Vegetation and community types			冠层 Foliage
植被类型和群落类型 Vegetation and community types			碳 C (%)	氮 N (g·kg^-1)	磷 P (g·kg^-1)	碳 C (%)	氮 N (g·kg^-1)	磷 P (g·kg^-1)
常绿阔叶林 Evergreen broad-leaved forests
栲树群落﹟		栲树 Castanopsis fargesii	43.8 (1.6)	13.9 (1.5)	0.7 (0.1)	46.2 (1.4)	9.0 (1.6)	0.6 (0.1)
		狗骨柴 Tricalysia dubia	44.2 (2.2)	17.6 (2.1)	0.9 (0.1)	39.7 (1.7)	4.9 (1.4)	0.4 (0.1)
		黄丹木姜子 Litsea elongata	48.6 (2.5)	11.5 (1.7)	0.7 (0.1)	46.4 (1.9)	4.3 (0.6)	0.5 (0.1)
		连蕊茶 Camellia fraterna	41.9 (2.6)	10.4 (2.3)	0.5 (0.1)	42.4 (3.9)	4.9 (1.0)	0.4 (0.1)
		天竺桂 Cinnamomum japonicum	49.1 (1.4)	12.1 (1.7)	0.7 (0.1)	47.2 (1.8)	5.0 (0.9)	0.5 (0.1)
		交让木 Daphniphyllum macropodum	41.8 (2.2)	12.6 (1.6)	0.6 (0.1)	39.2 (2.1)	6.2 (1.1)	0.4 (0.1)
		笔罗子 Meliosma regida	43.2 (1.9)	9.4 (0.7)	0.7 (0.1)	39.7 (1.7)	4.9 (1.4)	0.5 (0.1)
木荷群落＊		木荷 Schima superba	49.7 (3.1)	9.7 (1.4)	0.6 (0.1)	44.9 (0.8)	7.7 (0.9)	0.5 (0.1)
		栲树 Castanopsis fargesii	46.2 (4.2)	13.6 (1.2)	0.6 (0.1)	42.1 (1.9)	11.2 (1.8)	0.7 (0.1)
		马尾松 Pinus massoniana	51.4 (1.2)	11.8 (1.5)	0.5 (0.1)	47.9 (0.4)	8.8 (0.1)	0.8 (0.1)
		石栎 Lithocarpus glaber	47.9 (1.2)	9.7 (1.3)	0.7 (0.1)	46.9 (5.2)	8.8 (1.1)	0.6 (0.1)
		连蕊茶 Camellia fraterna	43.5 (1.1)	10.7 (1.9)	0.7 (0.1)	40.4 (1.7)	7.4 (1.2)	0.7 (0.1)
		山矾 Symplocos sumuntia	32.0 (1.6)	9.3 (0.9)	0.5 (0.1)	33.5 (1.6)	6.0 (0.6)	0.5 (0.1)
		柃木属植物 Eurya spp.	40.5 (1.6)	8.9 (1.8)	0.4 (0.1)	37.5 (2.8)	6.6 (0.3)	0.7 (0.2)
常绿针叶林 Evergreen coniferous forests
马尾松木荷群落※	马尾松 Pinus massoniana		42.4 (5.7)	11.3 (0.6)	1.1 (0.1)	50.4 (1.7)	5.4 (0.4)	0.6 (0.1)
	木荷 Schima superba		33.6 (5.9)	9.1 (0.7)	0.5 (0.1)	47.7 (1.8)	6.4 (0.8)	0.5 (0.1)
	苦槠 Castanopsis sclerophylla		42.0 (5.0)	10.4 (1.2)	0.9 (0.1)	46.9 (1.3)	10.1 (0.7)	0.7 (0.1)
	栲树 Castanopsis fargesii		39.3 (1.0)	12.5 (1.2)	0.8 (0.2)	45.7 (1.2)	8.7 (1.2)	0.7 (0.1)
	赤皮青冈 Cyclobalanopsis gilva		35.5 (5.1)	14.6 (1.1)	0.9 (0.1)	46.6 (1.5)	10.4 (2.3)	0.7 (0.1)
	连蕊茶 Camellia fraterna		37.7 (5.9)	7.9 (1.7)	0.6 (0.1)	39.1 (1.7)	9.4 (0.5)	0.7 (0.1)
	披针叶山矾 Symplocos lancifolia		39.1 (5.6)	11.9 (1.3)	0.6 (0.1)	35.3 (1.6)	10.5 (1.5)	0.6 (0.1)
	山矾 Symplocos sumuntia		38.6 (3.7)	10.5 (1.9)	0.6 (0.1)	39.8 (3.6)	11.2 (0.9)	0.7 (0.1)
	窄基红褐柃 Eurya rubiginosa		36.0 (1.2)	11.0 (1.1)	0.7 (0.1)	42.3 (3.1)	7.2 (1.1)	0.5 (0.1)
马尾松群落§	马尾松 Pinus massoniana		51.4 (1.6)	5.8 (1.0)	0.5 (0.1)	48.7 (1.7)	5.4 (0.6)	0.5 (0.1)
	木荷 Schima superba		45.9 (1.2)	9.1 (1.3)	0.6 (0.1)	46.6 (1.3)	6.6 (0.6)	0.5 (0.1)
	苦槠 Castanopsis sclerophylla		46.1 (1.7)	8.5 (1.1)	0.5 (0.1)	45.5 (1.7)	9.3 (0.8)	0.7 (0.1)
	石栎 Lithocarpus glaber		51.6 (2.6)	8.1 (1.5)	0.6 (0.1)	43.0 (2.1)	10.9 (1.2)	0.6 (0.1)
	栲树 Castanopsis fargesii		47.4 (1.9)	8.3 (1.1)	0.6 (0.1)	43.2 (1.7)	10.4 (0.8)	0.8 (0.1)
	山矾 Symplocos sumuntia		32.9 (1.9)	9.5 (2.0)	0.6 (0.1)	31.7 (1.5)	10.3 (1.2)	0.7 (0.1)
	檵木 Loropetalum chinense		44.2 (1.6)	11.6 (1.5)	0.8 (0.1)	40.2 (4.3)	8.5 (2.3)	0.7 (0.1)
落叶阔叶林 Deciduous broad-leaved forests
枫香群落^∆	枫香 Liquidambar formosana		43.4 (1.8)	18.5 (2.0)	1.4 (0.1)	37.9 (1.3)	13.3 (1.1)	1.0 (0.1)
	化香 Platycarya strobilacea		43.6 (1.2)	16.7 (2.5)	1.8 (0.2)	35.5 (3.4)	15.4 (1.1)	1.2 (0.2)
	白栎 Quercus fabri		43.9 (1.4)	17.3 (1.5)	1.7 (0.1)	39.6 (2.0)	12.5 (1.7)	1.0 (0.2)
	青灰叶下珠 Phyllanthus glaucus		39.5 (1.3)	23.5 (3.5)	2.1 (0.5)	40.1 (3.6)	14.1 (1.7)	0.9 (0.2)
	天仙果 Ficus erecta var. beecheyana		36.6 (1.2)	18.3 (1.7)	1.3 (0.1)	39.2 (2.7)	13.8 (1.6)	0.9 (0.1)
	箬竹 Indocalamus tessellates		41.4 (1.1)	15.6 (1.2)	1.8 (0.2)	33.3 (3.5)	13.2 (1.9)	0.9 (0.1)
南酸枣群落☆	南酸枣 Choerospondias axillaries		36.3 (1.2)	14.4 (2.3)	1.8 (0.3)	44.0 (1.5)	7.9 (0.4)	0.5 (0.1)
	长叶石栎 Lithocarpus harlandii		40.0 (1.1)	11.5 (0.9)	1.4 (0.3)	46.9 (3.4)	8.4 (1.6)	0.5 (0.1)
	华东楠 Machilus leptophylla		41.5 (2.6)	12.5 (1.3)	1.1 (0.2)	42.1 (2.7)	11.6 (1.2)	0.8 (0.1)
	披针叶茴香 Illicium lanceolatum		38.5 (2.1)	8.3 (0.7)	0.9 (0.1)	45.6 (1.9)	7.6 (0.9)	0.5 (0.1)
	紫楠 Phoebe sheareri		39.7 (2.2)	14.3 (1.8)	1.0 (0.2)	38.7 (2.3)	8.8 (1.5)	0.6 (0.2)
	黄丹木姜子 Litsea elongata		37.6 (1.5)	11.6 (1.6)	0.8 (0.1)	45.9 (3.6)	6.6 (0.1)	0.6 (0.1)
	连蕊茶 Camellia fraterna		40.1 (1.3)	7.9 (0.5)	0.7 (0.1)	35.7 (4.5)	10.6 (0.6)	0.9 (0.1)
	赤杨叶 Alniphyllum fortunei		41.6 (1.4)	16.3 (1.6)	0.9 (0.1)	42.1 (4.4)	10.9 (1.1)	1.1 (0.1)

植被类型和群落类型 Vegetation and community types			冠层 Foliage
植被类型和群落类型 Vegetation and community types			碳 C (%)	氮 N (g·kg^-1)	磷 P (g·kg^-1)	碳 C (%)	氮 N (g·kg^-1)	磷 P (g·kg^-1)
常绿阔叶林 Evergreen broad-leaved forests
栲树群落﹟		栲树 Castanopsis fargesii	43.8 (1.6)	13.9 (1.5)	0.7 (0.1)	46.2 (1.4)	9.0 (1.6)	0.6 (0.1)
		狗骨柴 Tricalysia dubia	44.2 (2.2)	17.6 (2.1)	0.9 (0.1)	39.7 (1.7)	4.9 (1.4)	0.4 (0.1)
		黄丹木姜子 Litsea elongata	48.6 (2.5)	11.5 (1.7)	0.7 (0.1)	46.4 (1.9)	4.3 (0.6)	0.5 (0.1)
		连蕊茶 Camellia fraterna	41.9 (2.6)	10.4 (2.3)	0.5 (0.1)	42.4 (3.9)	4.9 (1.0)	0.4 (0.1)
		天竺桂 Cinnamomum japonicum	49.1 (1.4)	12.1 (1.7)	0.7 (0.1)	47.2 (1.8)	5.0 (0.9)	0.5 (0.1)
		交让木 Daphniphyllum macropodum	41.8 (2.2)	12.6 (1.6)	0.6 (0.1)	39.2 (2.1)	6.2 (1.1)	0.4 (0.1)
		笔罗子 Meliosma regida	43.2 (1.9)	9.4 (0.7)	0.7 (0.1)	39.7 (1.7)	4.9 (1.4)	0.5 (0.1)
木荷群落＊		木荷 Schima superba	49.7 (3.1)	9.7 (1.4)	0.6 (0.1)	44.9 (0.8)	7.7 (0.9)	0.5 (0.1)
		栲树 Castanopsis fargesii	46.2 (4.2)	13.6 (1.2)	0.6 (0.1)	42.1 (1.9)	11.2 (1.8)	0.7 (0.1)
		马尾松 Pinus massoniana	51.4 (1.2)	11.8 (1.5)	0.5 (0.1)	47.9 (0.4)	8.8 (0.1)	0.8 (0.1)
		石栎 Lithocarpus glaber	47.9 (1.2)	9.7 (1.3)	0.7 (0.1)	46.9 (5.2)	8.8 (1.1)	0.6 (0.1)
		连蕊茶 Camellia fraterna	43.5 (1.1)	10.7 (1.9)	0.7 (0.1)	40.4 (1.7)	7.4 (1.2)	0.7 (0.1)
		山矾 Symplocos sumuntia	32.0 (1.6)	9.3 (0.9)	0.5 (0.1)	33.5 (1.6)	6.0 (0.6)	0.5 (0.1)
		柃木属植物 Eurya spp.	40.5 (1.6)	8.9 (1.8)	0.4 (0.1)	37.5 (2.8)	6.6 (0.3)	0.7 (0.2)
常绿针叶林 Evergreen coniferous forests
马尾松木荷群落※	马尾松 Pinus massoniana		42.4 (5.7)	11.3 (0.6)	1.1 (0.1)	50.4 (1.7)	5.4 (0.4)	0.6 (0.1)
	木荷 Schima superba		33.6 (5.9)	9.1 (0.7)	0.5 (0.1)	47.7 (1.8)	6.4 (0.8)	0.5 (0.1)
	苦槠 Castanopsis sclerophylla		42.0 (5.0)	10.4 (1.2)	0.9 (0.1)	46.9 (1.3)	10.1 (0.7)	0.7 (0.1)
	栲树 Castanopsis fargesii		39.3 (1.0)	12.5 (1.2)	0.8 (0.2)	45.7 (1.2)	8.7 (1.2)	0.7 (0.1)
	赤皮青冈 Cyclobalanopsis gilva		35.5 (5.1)	14.6 (1.1)	0.9 (0.1)	46.6 (1.5)	10.4 (2.3)	0.7 (0.1)
	连蕊茶 Camellia fraterna		37.7 (5.9)	7.9 (1.7)	0.6 (0.1)	39.1 (1.7)	9.4 (0.5)	0.7 (0.1)
	披针叶山矾 Symplocos lancifolia		39.1 (5.6)	11.9 (1.3)	0.6 (0.1)	35.3 (1.6)	10.5 (1.5)	0.6 (0.1)
	山矾 Symplocos sumuntia		38.6 (3.7)	10.5 (1.9)	0.6 (0.1)	39.8 (3.6)	11.2 (0.9)	0.7 (0.1)
	窄基红褐柃 Eurya rubiginosa		36.0 (1.2)	11.0 (1.1)	0.7 (0.1)	42.3 (3.1)	7.2 (1.1)	0.5 (0.1)
马尾松群落§	马尾松 Pinus massoniana		51.4 (1.6)	5.8 (1.0)	0.5 (0.1)	48.7 (1.7)	5.4 (0.6)	0.5 (0.1)
	木荷 Schima superba		45.9 (1.2)	9.1 (1.3)	0.6 (0.1)	46.6 (1.3)	6.6 (0.6)	0.5 (0.1)
	苦槠 Castanopsis sclerophylla		46.1 (1.7)	8.5 (1.1)	0.5 (0.1)	45.5 (1.7)	9.3 (0.8)	0.7 (0.1)
	石栎 Lithocarpus glaber		51.6 (2.6)	8.1 (1.5)	0.6 (0.1)	43.0 (2.1)	10.9 (1.2)	0.6 (0.1)
	栲树 Castanopsis fargesii		47.4 (1.9)	8.3 (1.1)	0.6 (0.1)	43.2 (1.7)	10.4 (0.8)	0.8 (0.1)
	山矾 Symplocos sumuntia		32.9 (1.9)	9.5 (2.0)	0.6 (0.1)	31.7 (1.5)	10.3 (1.2)	0.7 (0.1)
	檵木 Loropetalum chinense		44.2 (1.6)	11.6 (1.5)	0.8 (0.1)	40.2 (4.3)	8.5 (2.3)	0.7 (0.1)
落叶阔叶林 Deciduous broad-leaved forests
枫香群落^∆	枫香 Liquidambar formosana		43.4 (1.8)	18.5 (2.0)	1.4 (0.1)	37.9 (1.3)	13.3 (1.1)	1.0 (0.1)
	化香 Platycarya strobilacea		43.6 (1.2)	16.7 (2.5)	1.8 (0.2)	35.5 (3.4)	15.4 (1.1)	1.2 (0.2)
	白栎 Quercus fabri		43.9 (1.4)	17.3 (1.5)	1.7 (0.1)	39.6 (2.0)	12.5 (1.7)	1.0 (0.2)
	青灰叶下珠 Phyllanthus glaucus		39.5 (1.3)	23.5 (3.5)	2.1 (0.5)	40.1 (3.6)	14.1 (1.7)	0.9 (0.2)
	天仙果 Ficus erecta var. beecheyana		36.6 (1.2)	18.3 (1.7)	1.3 (0.1)	39.2 (2.7)	13.8 (1.6)	0.9 (0.1)
	箬竹 Indocalamus tessellates		41.4 (1.1)	15.6 (1.2)	1.8 (0.2)	33.3 (3.5)	13.2 (1.9)	0.9 (0.1)
南酸枣群落☆	南酸枣 Choerospondias axillaries		36.3 (1.2)	14.4 (2.3)	1.8 (0.3)	44.0 (1.5)	7.9 (0.4)	0.5 (0.1)
	长叶石栎 Lithocarpus harlandii		40.0 (1.1)	11.5 (0.9)	1.4 (0.3)	46.9 (3.4)	8.4 (1.6)	0.5 (0.1)
	华东楠 Machilus leptophylla		41.5 (2.6)	12.5 (1.3)	1.1 (0.2)	42.1 (2.7)	11.6 (1.2)	0.8 (0.1)
	披针叶茴香 Illicium lanceolatum		38.5 (2.1)	8.3 (0.7)	0.9 (0.1)	45.6 (1.9)	7.6 (0.9)	0.5 (0.1)
	紫楠 Phoebe sheareri		39.7 (2.2)	14.3 (1.8)	1.0 (0.2)	38.7 (2.3)	8.8 (1.5)	0.6 (0.2)
	黄丹木姜子 Litsea elongata		37.6 (1.5)	11.6 (1.6)	0.8 (0.1)	45.9 (3.6)	6.6 (0.1)	0.6 (0.1)
	连蕊茶 Camellia fraterna		40.1 (1.3)	7.9 (0.5)	0.7 (0.1)	35.7 (4.5)	10.6 (0.6)	0.9 (0.1)
	赤杨叶 Alniphyllum fortunei		41.6 (1.4)	16.3 (1.6)	0.9 (0.1)	42.1 (4.4)	10.9 (1.1)	1.1 (0.1)

层次 Layer	总体 Overall		常绿阔叶林 Evergreen broad-leaved forests		常绿针叶林 Evergreen coniferous forests		落叶阔叶林 Deciduous broad-leaved forests
层次 Layer	b (conf. inter.)	R²(n)	b (conf. inter.)	R²(n)	b (conf. inter.)	R²(n)	b (conf. inter.)	R²(n)
冠层 Foliage
乔木 Tree	1.39 (1.15-1.69)	0.47 (58)	1.03^* (0.65-1.64)	0.31 (16)	1.04 (0.8-1.35)	0.58 (27)	1.03^** (0.66-1.59)	0.44 (15)
灌木 Shrub	1.21 (1.05-1.39)	0.62 (78)	1.0 (0.77-1.31)	0.57 (27)	0.67 (0.5-0.9)	0.6 (21)	1.08 (0.88-1.32)	0.72 (30)
整体 Total	1.29 (1.15-1.45)	0.54 (136)	1.01 (0.81-1.27)	0.47 (43)	0.92 (0.76-1.13)	0.54 (48)	1.12 (0.93-1.35)	0.63 (45)
凋落物 Senesced litter
乔木 Tree	0.92 (0.76-1.1)	0.53 (58)			0.73 (0.53-0.99)	0.41 (27)	1.44 (1.29-1.6)	0.97 (15)
灌木 Shrub	0.89 (0.77-1.04)	0.53 (78)	0.75^** (0.54-1.04)	0.34 (27)	0.7 (0.51-0.95)	0.57 (21)	0.99 (0.8-1.24)	0.68 (30)
整体 Total	0.9 (0.8-1.01)	0.53 (136)	0.76 (0.58-0.98)	0.31 (43)	0.69 (0.56-0.86)	0.49 (48)	1.09 (0.92-1.28)	0.72 (45)

层次 Layer	总体 Overall		常绿阔叶林 Evergreen broad-leaved forests		常绿针叶林 Evergreen coniferous forests		落叶阔叶林 Deciduous broad-leaved forests
层次 Layer	b (conf. inter.)	R²(n)	b (conf. inter.)	R²(n)	b (conf. inter.)	R²(n)	b (conf. inter.)	R²(n)
冠层 Foliage
乔木 Tree	1.39 (1.15-1.69)	0.47 (58)	1.03^* (0.65-1.64)	0.31 (16)	1.04 (0.8-1.35)	0.58 (27)	1.03^** (0.66-1.59)	0.44 (15)
灌木 Shrub	1.21 (1.05-1.39)	0.62 (78)	1.0 (0.77-1.31)	0.57 (27)	0.67 (0.5-0.9)	0.6 (21)	1.08 (0.88-1.32)	0.72 (30)
整体 Total	1.29 (1.15-1.45)	0.54 (136)	1.01 (0.81-1.27)	0.47 (43)	0.92 (0.76-1.13)	0.54 (48)	1.12 (0.93-1.35)	0.63 (45)
凋落物 Senesced litter
乔木 Tree	0.92 (0.76-1.1)	0.53 (58)			0.73 (0.53-0.99)	0.41 (27)	1.44 (1.29-1.6)	0.97 (15)
灌木 Shrub	0.89 (0.77-1.04)	0.53 (78)	0.75^** (0.54-1.04)	0.34 (27)	0.7 (0.51-0.95)	0.57 (21)	0.99 (0.8-1.24)	0.68 (30)
整体 Total	0.9 (0.8-1.01)	0.53 (136)	0.76 (0.58-0.98)	0.31 (43)	0.69 (0.56-0.86)	0.49 (48)	1.09 (0.92-1.28)	0.72 (45)