Analysis of grassland community-level plant functional traits and functional diversity at different times during restoration

doi:10.11686/cyxb2022029

Abstract

Abstract:

Plant functional traits and functional diversity are important factors when evaluating the function of grassland ecosystems. In this study， we conducted a series of analyses of the understory grassland of Robinia pseudoacacia forests and abandoned grassland at different time points during restoration in the hilly and gully regions of the Loess Plateau. The community-level dynamics of plant functional traits and functional diversity were analyzed during vegetation restoration to explore how the grassland plant community and its functional traits change over time. The results showed that： 1） The understory grassland and abandoned grassland were both dominated by perennial weeds in the Asteraceae and Rosaceae； 2） During restoration， the understory grassland succeeded from a dicot-dominated community to a monocot-dominated community， while Asteraceae species were gradually replaced by Poaceae species. The abandoned grassland was always dominated by Asteraceae species， however， and showed a tendency to shrub encroachment. 3） The values of community-level leaf functional traits were higher in understory grassland than in abandoned grassland， while the values of community-level plant functional traits of coverage， density， and aboveground biomass were lower in understory grassland than in abandoned grassland. The values of species diversity and functional trait diversity were higher in abandoned grassland than in understory grassland， and abandoned grassland exhibited higher ecological benefits. These results provide a theoretical basis for the evaluation of grassland ecosystem functions in the hill and gully region of the Loess Plateau.

Key words: Loess Plateau, restoration grassland, community-level traits, functional diversity, species composition

Jiang-wen LI, Bang-yin HE, Cai LI, Hong-yan HUI, Bo LIU, Xiao-xi ZHANG, Hui FAN, Wen-yu SU. Analysis of grassland community-level plant functional traits and functional diversity at different times during restoration[J]. Acta Prataculturae Sinica, 2023, 32(1): 16-25.

Figures/Tables 6

References 39

1	Petchey O L， Gaston K J. Functional diversity （FD）， species richness and community composition. Ecology Letters， 2002， 5（3）： 402-411.
2	Petchey O L， Gaston K J. Functional diversity： Back to basics and looking forward. Ecology Letters， 2006， 9（6）： 741-758.
3	Xiang X， Huang Y M， Yang C Y， et al. Effect of altitude on community-level plant functional traits in the Qinghai Lake Basin， China. Chinese Journal of Plant Ecology， 2021， 45（5）： 456-466.
	向响，黄永梅，杨崇曜，等. 海拔对青海湖流域群落水平植物功能性状的影响. 植物生态学报， 2021， 45（5）： 456-466.
4	Liu J， Wu J， Su H， et al. Effects of grazing exclusion in Xilin Gol grassland differ between regions. Ecological Engineering， 2017， 99： 271-281.
5	Chai Q L. Effects of different managements on community structure and ecological functions in a typical steppe. Xianyang： Northwest A&F University， 2019.
	柴清琳. 不同管理措施对典型草原群落结构和生态功能的影响. 咸阳：西北农林科技大学， 2019.
6	Ding W， Wang Y B， Xiang G H， et al. Effects of Caragana microphylla encroachment on community structure and ecosystem function of a typical steppe. Chinese Journal of Plant Ecology， 2020， 44（1）： 33-43.
	丁威，王玉冰，向官海，等. 小叶锦鸡儿灌丛化对典型草原群落结构与生态系统功能的影响. 植物生态学报， 2020， 44（1）： 33-43.
7	Guo P， Xie L N， Man L， et al. Effects of the expansion of Caragana shrubs on forage yield and plant diversity in desert steppe. Pratacultural Science， 2019， 36（5）： 1215-1223.
	郭璞，解李娜，满良，等. 荒漠化草原锦鸡儿属灌丛扩增对牧草产量和植物多样性的影响. 草业科学， 2019， 36（5）： 1215-1223.
8	Qu H， Wang B T， Wang D， et al. Research of artificial forest plant diversity under different configuration of loess area. Ecology and Environmental Sciences， 2010， 19（4）： 843-848.
	曲红，王百田，王棣，等. 黄土区不同配置人工林物种多样性研究. 生态环境学报， 2010， 19（4）： 843-848.
9	Yu X， Wang X， Wu T， et al. Effects of enclosure on species diversity and functional diversity of desert steppe. Journal of Soil and Water Conservation， 2021， 35（6）： 243-250.
	余轩，王兴，吴婷，等. 围封对荒漠草原物种多样性和功能多样性的影响. 水土保持学报， 2021， 35（6）： 243-250.
10	Shannon C， Weaver W. The mathematical theory of communication. Urbana-USA： University of Illinois Press， 1949.
11	Pielou E C. Ecological diversity. New York： Wiley， 1975.
12	Simpson E H. Measurements of diversity. Nature， 1949（163）： 688.
13	Mason N W H， De-bello F. Functional diversity： A tool for answering challenging ecological questions. Journal of Vegetation Science， 2013， 24（5）： 777-780.
14	Mason N W H， Mac-Gillivray K， Steel J B， et al. An index of functional diversity. Journal of Vegetation Science， 2003， 14： 571-578.
15	Casanoves F， Pla L， Rienzo J A D， et al. FDiversity： A software package for the integrated analysis of functional diversity. Methods in Ecology and Evolution， 2011， 2（3）： 233-237.
16	Villéger S， Mason N W， Mouillot D. New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology， 2008， 89（8）： 2290-2301.
17	Mason N W H， De-bello F， Mouillot D， et al. A guide for using functional diversity indices to reveal changes in assembly processes along ecological gradients. Journal of Vegetation Science， 2013， 24（5）： 794-806.
18	Wang S M， Pang Y X， Song A Y， et al. Soil physiochemical properties and diversity of herbaceous plants dynamic on the different ages mixed forests of Populus×Euramercana ‘Neva’ and Robinia pseucdoacacia in coastal saline-alkali area. Acta Ecologica Sinica， 2018， 38（18）： 6539-6548.
	王树梅，庞元湘，宋爱云，等. 基于林龄的滨海盐碱地杨树刺槐混交林土壤理化性质及草本植物多样性动态. 生态学报， 2018， 38（18）： 6539-6548.
19	Liu J L， Yang Z L， Dang P， et al. Response of soil microbial community dynamics to Robinia pseudoacacia L. afforestation in the loess plateau： A chronosequence approach. Plant and Soil， 2017， 423（1）： 327-338.
20	Wang M Z， Bi H J， Jin S， et al. Effects of stand density on understory species diversity and soil physicochemical properties of a Cupressus funebris plantation in Yunding Mountain. Acta Ecologica Sinica， 2019， 39（3）： 981-988．
	王媚臻，毕浩杰，金锁，等. 林分密度对云顶山柏木人工林林下物种多样性和土壤理化性质的影响. 生态学报， 2019， 39（3）： 981-988.
21	Zhang W， Liu W C， Xu M P， et al. Response of forest growth to C∶N∶P stoichiometry in plants and soils during Robinia pseudoacacia afforestation on the Loess Plateau， China. Geoderma， 2019， 337： 280-289.
22	Wang J， Zhao W W， Zhang X， et al. Effects of reforestation on plant species diversity on the Loess Plateau of China： A case study in Danangou catchment. Science of the Total Environment， 2019， 651： 979-989.
23	Qiao W J， Dai Y Y， Zhang W， et al. Relationship between the vegetation community and soil nutrient and enzyme activity during the restoration of abandoned land in the Loess Hilly Region. Environmental Science， 2018， 39（12）： 5687-5698.
	乔文静，戴银月，张伟，等. 黄土丘陵区撂荒恢复过程中植物群落组成与土壤养分及酶活性变化的关系. 环境科学， 2018， 39（12）： 5687-5698.
24	Yang L X， Chen S F， An J J， et al. Relationships among community diversity and soil organic matter， total nitrogen under different vegetation types in the gully region of loess region. Acta Agrestia Sinica， 2014， 22（2）： 291-298.
	杨丽霞，陈少锋，安娟娟，等. 陕北黄土丘陵区不同植被类型群落多样性与土壤有机质、全氮关系研究. 草地学报， 2014， 22（2）： 291-298.
25	An Q Q， Qiao W Y， Li W J， et al. Effect of shrub encroachment on grassland community structure and above-ground biomass on the Loess Plateau. Acta Botanica Boreali-Occidentalia Sinica， 2021， 41（4）： 664-671.
	安琪琪，乔文英，李维军，等. 灌丛化对黄土高原草地植物群落结构和地上生物量的影响. 西北植物学报， 2021， 41（4）： 664-671.
26	Garnier E， Cortez J， Billès G， et al. Plant functional markers capture ecosystem properties during secondary succession. Ecology， 2004， 85： 2630-2637.
27	Díaz S， Lavorel S， De-bello F， et al. Incorporating plant functional diversity effects in ecosystem service assessments. Proceedings of the National Academy of Sciences of the United States of America， 2004， 104： 20684-20689.
28	Violle C， Reich P B， Pacala S W， et al. The emergence and promise of functional biogeography. Proceedings of the National Academy of Sciences of the United States of America， 2014， 111： 13690-13696.
29	Nunes A， Köbel M， Pinho P， et al. Which plant traits respond to aridity？ A critical step to assess functional diversity in Mediterranean drylands. Agricultural and Forest Meteorology， 2017， 239： 176-184.
30	Mitchell R M， Ames G M， Wright J P. Intra specific trait variability shapes leaf trait response to altered fire regimes. Annals of Botany， 2021， 127（4）： 543-552.
31	Ren C J， Zhao F Z， Kang D， et al. Linkages of C∶N∶P stoichiometry and bacterial community in soil following afforestation of former farmland. Forest Ecology and Management， 2016， 376： 59-66.
32	Fanin N， Bertrand I. Aboveground litter quality is a better predictor than belowground microbial communities when estimating carbon mineralization along a land-use gradient. Soil Biology and Biochemistry， 2016， 94（1）： 46-60.
33	Barliza J C， Pelaez J D L， Campo J. Recovery of biogeochemical processes in restored tropical dry forest on a coal mine spoil in La Guajira， Colombia. Land Degradation & Development， 2018， 29（9）： 3174-3183.
34	Alon M， Sternberg M. Effects of extreme drought on primary production， species composition and species diversity of a Mediterranean annual plant community. Journal of Vegetation Science， 2019， 30（6）： 1045-1055.
35	Li X G， Zhu Z H， Zhou X S， et al. Effects of clipping， fertilizing and watering on the relationship between species diversity， functional diversity and primary productivity in alpine meadow of China. Chinese Journal of Plant Ecology， 2011， 35（11）： 1136-1147.
	李晓刚，朱志红，周晓松，等. 刈割、施肥和浇水对高寒草甸物种多样性与初级生产力关系的影响. 植物生态学报， 2011， 35（11）： 1136-1147.
36	Ge Z X， Sun G L， Yuan Y， et al. Herbaceous plant species diversity and functional diversity in the forest-steppe zone of Hebei， China. Acta Prataculturae Sinica， 2017， 26（7）： 35-44.
	葛兆轩，孙国龙，袁业，等. 河北省森林草原区草本植物物种多样性和功能多样性. 草业学报， 2017， 26（7）： 35-44.
37	Guo M M， Wang W L， Kang H L， et al. Changes in soil properties and erodibility of gully heads induced by vegetation restoration on the Loess Plateau， China. Journal of Arid Land， 2018， 10（5）： 712-725.
38	Kong D X， Miao C Y， Borthwick A G L， et al. Spatiotemporal variations in vegetation cover on the Loess Plateau， China， between 1982 and 2013： Possible causes and potential impacts. Environmental Science and Pollution Research， 2018， 25（14）： 13633-13644.
39	Tuo D F， Gao G Y， Chang R Y， et al. Effects of revegetation and precipitation gradient on soil carbon and nitrogen variations in deep profiles on the Loess Plateau of China. Science of the Total Environment， 2018， 626： 399-411.

恢复年限 Restoration years （a）	平均海拔 Altitude （m）	坡向 Aspect	林下草地Understory grassland		撂荒草地Abandoned grassland
恢复年限 Restoration years （a）	平均海拔 Altitude （m）	坡向 Aspect	坐标Coordinate	主要物种Main species	坐标Coordinate	主要物种Main species
10	1337	东南Southeast	N 36°44′05″， E 109°15′14″	猪毛蒿A. scoparia、山莓Rubus corchorifolius	N 36°44′06″， E 109°15′13″	阿尔泰狗娃花Heteropappus altaicus、狗尾草S. viridis
20	1208	正南South	N 36°43′44″， E 109°15′13″	野菊C. indicum、异叶败酱Patrinia heterophylla	N 36°43′40″， E 109°14′14″	小花鬼针草Bidens parviflora、早熟禾Poa annua
30	1155	东南Southeast	N 36°44′45″， E 109°15′18″	铁杆蒿A. sacrorum、龙牙草Agrimonia pilosa	N 36°44′43″， E 109°14′48″	猪毛蒿A. scoparia、二裂委陵菜Potentilla bifurca
45	1254	正南South	N 36°44′39″， E 109°15′30″	披碱草E. dahuricus、艾蒿Artemisia argyi	N 36°44′29″， E 109°15′26″	黄刺玫R. xanthina、苦荬菜Ixeris polycephala

恢复年限 Restoration years （a）	平均海拔 Altitude （m）	坡向 Aspect	林下草地Understory grassland		撂荒草地Abandoned grassland
恢复年限 Restoration years （a）	平均海拔 Altitude （m）	坡向 Aspect	坐标Coordinate	主要物种Main species	坐标Coordinate	主要物种Main species
10	1337	东南Southeast	N 36°44′05″， E 109°15′14″	猪毛蒿A. scoparia、山莓Rubus corchorifolius	N 36°44′06″， E 109°15′13″	阿尔泰狗娃花Heteropappus altaicus、狗尾草S. viridis
20	1208	正南South	N 36°43′44″， E 109°15′13″	野菊C. indicum、异叶败酱Patrinia heterophylla	N 36°43′40″， E 109°14′14″	小花鬼针草Bidens parviflora、早熟禾Poa annua
30	1155	东南Southeast	N 36°44′45″， E 109°15′18″	铁杆蒿A. sacrorum、龙牙草Agrimonia pilosa	N 36°44′43″， E 109°14′48″	猪毛蒿A. scoparia、二裂委陵菜Potentilla bifurca
45	1254	正南South	N 36°44′39″， E 109°15′30″	披碱草E. dahuricus、艾蒿Artemisia argyi	N 36°44′29″， E 109°15′26″	黄刺玫R. xanthina、苦荬菜Ixeris polycephala

群落加权平均性状 Community weighted mean trait	草地类型 Grassland types	10 a	20 a	30 a	45 a
CWM_高度 Average height of CWM	林下草地Understory grassland	41.04±2.08Aa	44.90±4.25Aa	39.43±4.77Aa	51.81±7.04Aa
CWM_高度 Average height of CWM	撂荒草地Abandoned grassland	39.41±1.78Aa	32.63±4.92Aa	36.40±6.71Aa	42.46±3.08Aa
CWM_盖度 Coverage of CWM	林下草地Understory grassland	15.83±6.84Aa	8.19±1.90Aa	22.42±4.40Aa	15.37±4.03Ba
CWM_盖度 Coverage of CWM	撂荒草地Abandoned grassland	16.18±2.73Aab	12.16±3.21Ab	25.88±7.16Aab	30.00±3.24Aa
CWM_密度 Density of CWM	林下草地Understory grassland	7.04±2.68Ba	17.22±1.81Aa	24.08±17.84Aa	21.09±5.43Ba
CWM_密度 Density of CWM	撂荒草地Abandoned grassland	106.20±36.24Aa	27.50±8.90Ab	24.29±6.65Ab	111.44±22.81Aa
CWM_{地上生物量} Aboveground biomass of CWM	林下草地Understory grassland	24.18±7.00Aab	16.45±3.30Ab	78.78±34.61Aa	27.18±4.47Bab
CWM_{地上生物量} Aboveground biomass of CWM	撂荒草地Abandoned grassland	24.38±4.22Ab	23.47±4.69Ab	54.87±20.05Aab	63.83±11.10Aa
CWM_叶面积 Leaf area of CWM	林下草地Understory grassland	2310.50±301.23Aa	413.26±87.09Ab	1175.51±425.70Ab	1097.77±226.14Ab
CWM_叶面积 Leaf area of CWM	撂荒草地Abandoned grassland	364.26±65.64Bb	510.23±87.97Aab	677.10±78.28Aa	101.14±18.41Bc
CWM_叶片质量 Leaf biomass of CWM	林下草地Understory grassland	0.10±0.01Aa	0.02±0.00Ab	0.06±0.02Aab	0.06±0.01Aab
CWM_叶片质量 Leaf biomass of CWM	撂荒草地Abandoned grassland	0.02±0.00Bbc	0.03±0.01Aab	0.04±0.01Aa	0.01±0.00Bc

群落加权平均性状 Community weighted mean trait	草地类型 Grassland types	10 a	20 a	30 a	45 a
CWM_高度 Average height of CWM	林下草地Understory grassland	41.04±2.08Aa	44.90±4.25Aa	39.43±4.77Aa	51.81±7.04Aa
CWM_高度 Average height of CWM	撂荒草地Abandoned grassland	39.41±1.78Aa	32.63±4.92Aa	36.40±6.71Aa	42.46±3.08Aa
CWM_盖度 Coverage of CWM	林下草地Understory grassland	15.83±6.84Aa	8.19±1.90Aa	22.42±4.40Aa	15.37±4.03Ba
CWM_盖度 Coverage of CWM	撂荒草地Abandoned grassland	16.18±2.73Aab	12.16±3.21Ab	25.88±7.16Aab	30.00±3.24Aa
CWM_密度 Density of CWM	林下草地Understory grassland	7.04±2.68Ba	17.22±1.81Aa	24.08±17.84Aa	21.09±5.43Ba
CWM_密度 Density of CWM	撂荒草地Abandoned grassland	106.20±36.24Aa	27.50±8.90Ab	24.29±6.65Ab	111.44±22.81Aa
CWM_{地上生物量} Aboveground biomass of CWM	林下草地Understory grassland	24.18±7.00Aab	16.45±3.30Ab	78.78±34.61Aa	27.18±4.47Bab
CWM_{地上生物量} Aboveground biomass of CWM	撂荒草地Abandoned grassland	24.38±4.22Ab	23.47±4.69Ab	54.87±20.05Aab	63.83±11.10Aa
CWM_叶面积 Leaf area of CWM	林下草地Understory grassland	2310.50±301.23Aa	413.26±87.09Ab	1175.51±425.70Ab	1097.77±226.14Ab
CWM_叶面积 Leaf area of CWM	撂荒草地Abandoned grassland	364.26±65.64Bb	510.23±87.97Aab	677.10±78.28Aa	101.14±18.41Bc
CWM_叶片质量 Leaf biomass of CWM	林下草地Understory grassland	0.10±0.01Aa	0.02±0.00Ab	0.06±0.02Aab	0.06±0.01Aab
CWM_叶片质量 Leaf biomass of CWM	撂荒草地Abandoned grassland	0.02±0.00Bbc	0.03±0.01Aab	0.04±0.01Aa	0.01±0.00Bc

[1]	Zhan-dong PAN, Qian-qian MA, Xiao-long CHEN, Li-qun CAI, Xue-mei CAI, Bo DONG, Jun WU, Ren-zhi ZHANG. Effects of biochar addition on nutrient levels and humus and its components in dry farmland soils on the Loess Plateau [J]. Acta Prataculturae Sinica, 2022, 31(2): 14-24.
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