Effects of biodiversity on primary productivity and its mechanism in artificially sown clonal plant communities of the Sanjiangyuan region

doi:10.11686/cyxb2022446

Abstract

Abstract:

Clonal plants have unique ways to compete and reproduce， and their unique traits may lead to a particular biodiversity-productivity relationship in communities they form. However， this issue has rarely been studied. In this research， a biodiversity experiment was set up using three clonal species （Elymus nutans， Medicago ruthenica and Potentilla anserine） in monocultures and mixtures. There were three replicates， E. nutans was used as the fixed species in all mixtures and the effects of biodiversity on productivity were assessed over two years. We found that the net biodiversity effects （NE） were significant in nearly all mixtures in our experiment， attributable to strong trait-independent complementary effects （TICE）， although there was no positive linear relationship between species richness and aboveground biomass. The strong TICE was predominantly caused by the complementary utilization of aboveground vertical space， while a secondary cause was the enhancement of light absorption capacity of the understorey species P. anserine. Meanwhile， the dominance of E. nutans increased over time and with increasing species richness， whereas P. anserine displayed the opposite tendencies， resulting in an increase in trait-dependent complementary effect （TDCE） and dominance effect （DE） for communities where P. anserine was present. However， the contributions of those two mechanisms to NE were limited. Furthermore， all those effects increased with time. The results of this study indicate that clonal species display a different biodiversity-productivity relationship from the normally positive linear pattern， due to the complementary use of aboveground vertical space.

Key words: the biodiversity-productivity relationship, clonal plants, net biodiversity effects, trait-independent complementary effect, plant traits

Zeng-hui LIU, Su-jin LU, Yu-xin WANG, Chun-hui ZHANG, Xin YIN. Effects of biodiversity on primary productivity and its mechanism in artificially sown clonal plant communities of the Sanjiangyuan region[J]. Acta Prataculturae Sinica, 2023, 32(9): 27-38.

Figures/Tables 6

Fig.1 The effects of species composition and species richness on aboveground biomass in clonal plant community

Table 1 The effects of species richness and year on NE， TICE， TDCE and DE

处理 Treatment	df	NE		TICE		TDCE		DE
处理 Treatment	df	F	P	F	P	F	P	F	P
物种丰富度Species richness	1，14	2.058	0.1734	0.215	0.6497	1.583	0.22890	0.018	0.8945
年份Year	1，14	51.592	<0.0010	8.521	0.0112	14.385	0.00198	8.652	0.0107
丰富度×年份Species richness×year	1，14	6.057	0.0274	0.822	0.3798	1.934	0.18607	1.575	0.2300

Table 2 The effects of species composition and year on NE， TICE， TDCE and DE

处理 Treatment	df	NE		TICE		TDCE		DE
处理 Treatment	df	F	P	F	P	F	P	F	P
物种组成Species composition	1，14	1.867	0.197	0.416	0.6688	0.789	0.476	0.075	0.9286
年份Year	1，14	50.607	<0.001	7.700	0.0168	12.602	0.004	7.602	0.0174
物种组成×年份Species composition×year	1，14	2.980	0.089	0.378	0.6929	0.883	0.439	0.776	0.4819

Fig.2 The effects of species richness or species composition on net biodiversity effects （NE）， trait-independent complementary effects （TICE）， trait-dependent complementary effects （TDCE） and dominance effects （DE） in different years

Fig.3 The values of natural logarithm response ratio （LNRR） of aboveground biomass about different species in all mixture of different years

Fig.4 The effects of same plant species under different sowing mode （single and mixed sowing） on plant traits

References 56

1	Mori A S， Dee L E， Gonzalez A， et al. Biodiversity-productivity relationships are key to nature-based climate solutions. Nature Climate Change， 2021， 11（7）： 543-550.
2	He M， Pan Y， Zhou G， et al. Grazing and global change factors differentially affect biodiversity-ecosystem functioning relationships in grassland ecosystems. Global Change Biology， 2022， 28（18）： 5492-5504.
3	Xi N， Chen D， Bahn M， et al. Drought soil legacy alters drivers of plant diversity-productivity relationships in old-field systems. Science Advances， 2022， 8（18）： 1-11.
4	Tilman D. The ecological consequences of changes in biodiversity： A search for general principles. Ecology， 1999， 80（5）： 1455-1474.
5	Isbell F， Balvanera P， Mori A S， et al. Expert perspectives on global biodiversity loss and its drivers and impacts on people. Frontiers in Ecology and the Environment， 2022， 22（1）： 1-10.
6	Grime J P. Competition exclusion in herbaceous vegetation. Nature， 1973， 242： 344-347.
7	Gillman L N， Wright S D. The influence of productivity on the species richness of plants： A critical assessment. Ecology， 2006， 87（5）： 1234-1243.
8	Van R J， Berendse F. Diversity-productivity relationships： Initial effects， long-term patterns， and underlying mechanisms. Proceedings of the National Academy of Sciences of the United States of America， 2005， 102（3）： 695-700.
9	Roscher C， Schumacher J. Positive diversity effects on productivity in mixtures of arable weed species as related to density-size relationships. Journal of Plant Ecology， 2016， 9（6）： 792-804.
10	Cheng Y， Zhang C， Zhao X， et al. Biomass-dominant species shape the productivity-diversity relationship in two temperate forests. Annals of Forest Science， 2018， 75（4）： 97-106.
11	Chanteloup P， Bonis A. Functional diversity in root and above-ground traits in a fertile grassland shows a detrimental effect on productivity. Basic and Applied Ecology， 2013， 14（3）： 208-216.
12	Waide R B， Willig M R， Steiner C F， et al. The relationship between productivity and species richness. Annual Review of Ecology and Systematics， 1999， 30： 257-300.
13	Šímová I， Li Y M， Storch D. Relationship between species richness and productivity in plants： The role of sampling effect， heterogeneity and species pool. Journal of Ecology， 2013， 101（1）： 161-170.
14	Lisner A， Ottaviani G， Klimešová J， et al. The species richness-productivity relationship varies among regions and productivity estimates， but not with spatial resolution. Oikos， 2021， 130（10）： 1704-1714.
15	Long Z T， Bruno J F， Duffy J E. Biodiversity mediates productivity through different mechanisms at adjacent trophic levels. Ecology， 2007， 88（11）： 2821-2829.
16	Liu J J， Xu Y， Shan Y X， et al. Biotic and abiotic factors determine species diversity-productivity relationships in mountain meadows. Journal of Plant Ecology， 2021， 14（6）： 1175-1188.
17	Hillebrand H， Blasius B， Borer E T， et al. Biodiversity change is uncoupled from species richness trends： Consequences for conservation and monitoring. Journal of Applied Ecology， 2018， 55（1）： 169-184.
18	Spaak J W， Baert J M， Baird D J， et al. Shifts of community composition and population density substantially affect ecosystem function despite invariant richness. Ecology Letters， 2017， 20： 1315-1324.
19	Zhang R， Schellenberg M P， Tian D， et al. Shifting community composition determines the biodiversity-productivity relationship under increasing precipitation and N deposition. Journal of Vegetation Science， 2021， 32（2）： e12998.
20	Mahaut L， Fort F， Violle C， et al. Multiple facets of diversity effects on plant productivity： Species richness， functional diversity， species identity and intraspecific competition. Functional Ecology， 2020， 34（1）： 287-298.
21	Calatayud M L， Forrestel E J， Chang C C， et al. Demystifying dominant species. New Phytologist， 2019， 223（3）： 1106-1126.
22	Calatayud J， Andivia E， Escudero A， et al. Positive associations among rare species and their persistence in ecological assemblages. Nature Ecology & Evolution， 2020， 4（1）： 40-45.
23	Creed R P， Cherry R P， Pflaum J R， et al. Dominant species can produce a negative relationship between species diversity and ecosystem function. Oikos， 2009， 118（5）： 723-732.
24	O’Connor N E， Crowe T P. Biodiversity loss and ecosystem functioning： Distinguishing between number and identity of species. Ecology， 2005， 86（7）： 1783-1796.
25	Liancourt P D， Viard-Crétat F， Michalet R. Contrasting community responses to fertilization and the role of the competitive ability of dominant species. Journal of Vegetation Science， 2009， 20（1）： 138-147.
26	Hooper D U， Chapin F S， Ewel J J， et al. Effects of biodiversity on ecosystem functioning： A consensus of current knowledge. Ecological Monographs， 2005， 75（1）： 33-35.
27	Dong M， Yu F H， Peter A. Ecological consequences of plant clonality. Annals of Botany， 2014， 114（2）： 367.
28	Kang J J， Zhao W Z， Zhao M. Remediation of blowouts by clonal plants in Maqu degraded alpine grasslands of northwest China. Journal of Plant Research， 2017， 130（2）： 291-299.
29	Fu J J， Zhou H K， Zhao X Q， et al. Clonal plants and their importance of different alpine grasslands in Haibei region， Qinghai Province. Acta Agrestia Sinica， 2013， 21（6）： 1065-1072.
	付京晶，周华坤，赵新全，等. 青海海北不同类型高寒草地的克隆植物及其重要性. 草地学报， 2013， 21（6）： 1065-1072.
30	Loreau M， Hector A. Partitioning selection and complementarity in biodiversity experiments. Nature， 2001， 412（6842）： 72-76.
31	Fox J W. Interpreting the ‘selection effect’ of biodiversity on ecosystem function. Ecology Letters， 2005， 8（8）： 846-856.
32	Niklaus P A， Baruffol M， He J S， et al. Can niche plasticity promote biodiversity-productivity relationships through increased complementarity？ Ecology， 2017， 98（4）： 1104-1116.
33	Dong M. Effects of severing rhizome on clonal growth in rhizomatous grass species Psammochloa villosa and Leymus secalinus. Acta Botanica Sinica， 1999， 41（2）： 194-198.
	董鸣. 切断根茎对根茎禾草沙鞭和赖草克隆生长的影响. 植物学报， 1999， 41（2）： 194-198.
34	Wang J， Xu T， Wang Y， et al. A meta-analysis of effects of physiological integration in clonal plants under homogeneous vs. heterogeneous environments. Functional Ecology， 2021， 35（3）： 578-589.
35	Gilbert B， Turkington R， Srivastava D S. Dominant species and diversity： Linking relative abundance to controls of species establishment. The American Naturalist， 2009， 174（6）： 850-862.
36	Saiz H， Bittebiere A K， Benot M L， et al. Understanding clonal plant competition for space over time： A fine-scale spatial approach based on experimental communities. Journal of Vegetation Science， 2016， 27（4）： 759-770.
37	Hodapp D， Hillebrand H， Blasius B， et al. Environmental and trait variability constrain community structure and the biodiversity-productivity relationship. Ecology， 2016， 97（6）： 1463-1474.
38	Marquard E， Weigelt A， Temperton V M， et al. Plant species richness and functional composition drive overyielding in a six-year grassland experiment. Ecology， 2009， 90（12）： 3290-3302.
39	Kraft N J B， Godoy O， Levine J M. Plant functional traits and the multidimensional nature of species coexistence. Proceedings of the National Academy of Sciences of the United States of America， 2015， 112（3）： 797-802.
40	Gubsch M， Buchmann N， Schmid B， et al. Differential effects of plant diversity on functional trait variation of grass species. Annals of Botany， 2011， 107（1）： 157-169.
41	Vojtech E， Loreau M， Yachi S， et al. Light partitioning in experimental grass communities. Oikos， 2008， 117（9）： 1351-1361.
42	Flynn D F B， Mirotchnik N， Jain M， et al. Functional and phylogenetic diversity as predictors of biodiversity-ecosystem-functioning relationships. Ecology， 2011， 92： 1573-1581.
43	Gravel D， Bell T， Barbera C， et al. Experimental niche evolution alters the strength of the diversity-productivity relationship. Nature， 2011， 469（7328）： 89-94.
44	Zuppinger-Dingley D， Schmid B， Petermann J S， et al. Selection for niche differentiation in plant communities increases biodiversity effects. Nature， 2014， 515（7525）： 108-111.
45	Bao S D. Soil agrochemical analysis. Beijing： China Agriculture Press， 2000： 263-270.
	鲍士旦. 土壤农化分析. 北京：中国农业出版社， 2000： 263-270.
46	Wang L， Min J X， Shen H F， et al. R and its basic application in agricultural experiment analysis （Ⅱ）. China Rice， 2022， 28（1）： 95-102.
	王磊，闵佳鑫，申红芳，等. R语言及在农业试验数据分析中的基本应用（二）. 中国稻米， 2022， 28（1）： 95-102.
47	Van R J， Berendse F. Long-term persistence of a positive plant diversity-productivity relationship in the absence of legumes. Oikos， 2009， 118（1）： 101-106.
48	Tilman D， Reich P B， Knops J， et al. Diversity and productivity in a long-term grassland experiment. Science， 2001， 294（5543）： 843-845.
49	Barry K E， Kathryn E， Mommer L， et al. The future of complementarity： Disentangling causes from consequences. Trends in Ecology and Evolution， 2019， 34（2）： 167-180.
50	Bakker L M， Barry K E， Mommer L， et al. Focusing on individual plants to understand community scale biodiversity effects： the case of root distribution in grasslands. Oikos， 2021， 130（11）： 1954-1966.
51	Lorentzen S， Roscher C， Schumacher J， et al. Species richness and identity affect the use of aboveground space in experimental grasslands. Perspectives in Plant Ecology， Evolution and Systematics， 2008， 10（2）： 73-87.
52	Ishii H， Asano S. The role of crown architecture， leaf phenology and photosynthetic activity in promoting complementary use of light among coexisting species in temperate forests. Ecological Research， 2010， 25： 715-722.
53	Anten N P R， Hirose T. Shoot structure， leaf physiology， and daily carbon gain of plant species in a tallgrass meadow. Ecology， 2003， 84： 955-968.
54	Maestre F T， Ballini C， Baldy V， et al. On the relative importance of the effects of selection and complementarity as drivers of diversity productivity relationships in Mediterranean shrublands. Oikos， 2008， 117（9）： 1345-1350.
55	Yuan Z Q， Yu K L， Epstein H， et al. Effects of legume species introduction on vegetation and soil nutrient development on abandoned croplands in a semi-arid environment on the Loess Plateau， China. Science of the Total Environment， 2016， 541： 692-700.
56	Schöb C， Kerle S， Karley A J， et al. Intraspecific genetic diversity and composition modify species-level diversity-productivity relationships. New Phytologist， 2015， 205（2）： 720-730.