Fine-scale characteristics of the boundaries between annual patches and perennial patches in a meadow steppe

  • Yonghong Cao
  • Deli WangEmail author
  • Mikko Heino
  • Xincheng Li
  • Hui Zhu
  • Jushan Liu
  • Xuehui Zou
Research Article



Boundaries may have crucial influences on landscape patterns, processes, and dynamics. However, there is little understanding of mechanisms that govern changes in the location and composition of boundaries. At smaller scales, investigation of detailed soil and vegetation characteristics can clarify the linkages between soil properties and vegetation patterns.


The aims were to examine the relationship between vegetation patterns and soil properties, and to explore mechanisms that govern changes in the location and composition of boundaries.


In a 50-ha grassland fenced for more than 10 years, where a recovery process had been initiated and annual grasses dominated in most saline–alkaline areas, we quantitatively characterized the spatial gradients across the visually identified physiognomic boundary between annual patches and perennial patches at a fine spatial scale.


Fine-scale vegetation and soil boundaries were well-defined and statistically characterized by a high rate of change compared to immediately adjacent areas. Plant characteristics were markedly influenced by soil properties. The alteration of salinity and alkalinity were the most important factors explaining the plant patterns across patch boundaries. Successional processes of colonization were involved in perennial encroachment in the annual patches.


Underlying soil properties primarily determine the plant patterns of the boundary; plant succession caused by interspecific competition is superimposed on the plant–soil feedback loop maintaining soil nutrient conditions. These processes alter the characteristics and locations of patch boundaries in response to changing disturbance regimes. Our findings offer insight into how boundaries may respond to changes in environmental conditions and drive landscape-level dynamics.


Boundary characteristic Patch Meadow steppe Saline–alkaline grassland Fine-scale study Plant–soil feedback 



We thank László Körmöczi for helpful comments on an earlier draft of this paper. This project was supported by National Key Research and Development Program of China (2016YFC0500602), National Natural Science Foundation of China (No. 31770520), the Program for Introducing Talents to Universities (B16011), and the Program for Innovative Research Team in University (IRT-16R11).

Supplementary material

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Supplementary material 1 (XLSX 26 kb)
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Supplementary material 2 (DOCX 459 kb)


  1. Aguiar MR, Sala OE (1999) Patch structure, dynamics and implications for the functioning of arid ecosystems. Trends Ecol Evol 14:273–277CrossRefGoogle Scholar
  2. Allen CD, Breshears DD (1998) Drought-induced shift of a forest-woodland ecotone: rapid landscape response to climate variation. Proc Natl Acad Sci USA 95:14839–14842CrossRefGoogle Scholar
  3. Bai Z, Gao Y, Xing F, Sun S, Jiao D, Wei X, Mu C (2015) Responses of two contrasting saline–alkaline grassland communities to nitrogen addition during early secondary succession. J Veg Sci 26:686–696CrossRefGoogle Scholar
  4. Bestelmeyer BT, Ward JP, Havstad KM (2006) Soil-geomorphic heterogeneity governs patchy vegetation dynamics at an arid ecotone. Ecology 87:963–973CrossRefGoogle Scholar
  5. Buxbaum CAZ, Vanderbilt K (2007) Soil heterogeneity and the distribution of desert and steppe plant species across a desert-grassland ecotone. J Arid Environ 69:617–632CrossRefGoogle Scholar
  6. Carlile DW, Skalski JR, Batker JE, Thomas JM, Cullinan VI (1989) Determination of ecological scale. Landscape Ecol 2:203–213CrossRefGoogle Scholar
  7. Conti L, Bello F, Lepš J, Acosta ATR, Carboni M (2017) Environmental gradients and micro-heterogeneity shape fine-scale plant community assembly on coastal dunes. J Veg Sci 28:762–773CrossRefGoogle Scholar
  8. Corwin DL, Lesch SM (2005) Apparent soil electrical conductivity measurements in agriculture. Comput Electron Agric 46:11–43CrossRefGoogle Scholar
  9. Cramer W, Hytteborn H (1987) The separation of fluctuation and long-term change in vegetation dynamics of a rising seashore. Vegetatio 69:157–167CrossRefGoogle Scholar
  10. Dutoit T, Buisson E, Gerbaud E, Roche P, Tatoni T (2007) The status of transitions between cultivated fields and their boundaries: ecotones, ecoclines or edge effects? Acta Oecol 31:127–136CrossRefGoogle Scholar
  11. Fernandes MR, Segurado P, Jauch E, Ferreira MT (2016) Riparian responses to extreme climate and land-use change scenarios. Sci Total Environ 569–570:145–158CrossRefGoogle Scholar
  12. Ford H, Roberts A, Jones L (2016) Nitrogen and phosphorus co-limitation and grazing moderate nitrogen impacts on plant growth and nutrient cycling in sand dune grassland. Sci Total Environ Part A 542:203–209CrossRefGoogle Scholar
  13. Fortin MJ, Drapeau P, Jacquez GM (1996) Quantification of the spatial co-occurrences of ecological boundaries. Oikos 77:51–60CrossRefGoogle Scholar
  14. Gao Q, Li J, Zheng H (1996) A dynamic landscape simulation model for the alkaline grasslands on Songnen Plain in northeast China. Landscape Ecol 11:339–349CrossRefGoogle Scholar
  15. Han D, Yang Y, Yang Y, Li J, Yang Y (2012) Spatial patterns of plant species diversity in a degraded successional series of fragmented Leymus chinensis meadow in Songnen Plain of Northeast China. Chin J Appl Ecol 23:666–672Google Scholar
  16. Hobbs ER (1986) Characterizing the boundary between California annual grassland and coastal sage scrub with differential profiles. Plant Ecol 65:115–126CrossRefGoogle Scholar
  17. Hodgkinson KC (1998) Sprouting success of shrubs after fire: height-dependent relationships for different strategies. Oecologia 115:64–72CrossRefGoogle Scholar
  18. Ibrahim EA (2016) Seed priming to alleviate salinity stress in germinating seeds. J Plant Physiol 192:38–46CrossRefGoogle Scholar
  19. Jackson J (2005) Is there a relationship between herbaceous species richness and buffel grass (Cenchrus ciliaris)? Aust Ecol 30:505–517CrossRefGoogle Scholar
  20. Kindscher K, Wells PV (1995) Prairie plant guilds: a multivariate analysis of prairie species based on ecological and morphological traits. Vegetatio 117:29–50CrossRefGoogle Scholar
  21. Levin SA, Paine RT (1974) Disturbance, patch formation and community structure. Proc Natl Acad Sci USA 71:2744–2747CrossRefGoogle Scholar
  22. Li C, Fang B, Yang C, Shi D, Wang D (2009) Effects of various salt-alkaline mixed stresses on the state of mineral elements in nutrient solutions and the growth of alkali resistant halophyte Chloris virgata. J Plant Nutr 32:1137–1147CrossRefGoogle Scholar
  23. Liu J, Feng C, Wang D, Wang L, Wilsey BJ, Zhong Z (2015) Impacts of grazing by different large herbivores in grassland depend on plant species diversity. J Appl Ecol 52:1053–1062CrossRefGoogle Scholar
  24. Ludwig JA, Cornelius JM (1987) Locating discontinuities along ecological gradients. Ecology 68:448–450CrossRefGoogle Scholar
  25. Mason JA, Jacobs PM, Gruley KE, Reyerson P, Hanson PR (2016) Parent material influence on soil response to vegetation change, southeastern Minnesota, U.S.A. Geoderma 275:1–17CrossRefGoogle Scholar
  26. Montana C (1992) The colonization of bare areas in two-phase mosaics of an arid ecosystem. J Ecol 80:315–327CrossRefGoogle Scholar
  27. Moore KA (2009) Fluctuating patch boundaries in a native annual forb: the roles of niche and dispersal limitation. Ecology 90:378–387CrossRefGoogle Scholar
  28. Mouchet M, Guilhaumon F, Villger S, Manson NWH, Tomasini J-A, Mouillot D (2008) Towards a consensus for calculating dendrogram-based functional diversity indices. Oikos 117:794–800CrossRefGoogle Scholar
  29. Mulder CPH, Jumpponen A, Högberg P, Huss-Danell K (2002) How plant diversity and legumes affect nitrogen dynamics in experimental grassland communities. Oecologia 133:412–421CrossRefGoogle Scholar
  30. Müller SC, Overbeck GE, Pfadenhauer J, Pillar VD (2012) Woody species patterns at forest-grassland boundaries in southern Brazil. Flora 207:586–598CrossRefGoogle Scholar
  31. Muñoz-Reinoso JC (2009) Boundaries and scales in shrublands of the Doñana Biological Reserve, southwest Spain. Landscape Ecol 24:509–518CrossRefGoogle Scholar
  32. Nano CEM, Clarke PJ (2008) Variegated desert vegetation: covariation of edaphic and fire variables provides a framework for understanding mulga-spinifex coexistence. Aust Ecol 33:848–862CrossRefGoogle Scholar
  33. Nicholas AMM, Franklin DC, Bowman DMJS (2011) Floristic uniformity across abrupt boundaries between Triodia hummock grassland and Acacia shrubland on an Australian desert sandplain. J Arid Environ 75:1090–1096CrossRefGoogle Scholar
  34. Niu S, Liu W, Wan S (2008) Different growth responses of C3 and C4 grasses to seasonal water and nitrogen regimes and competition in a pot experiment. J Exp Bot 59:1431–1439CrossRefGoogle Scholar
  35. Oksanen J, Blanchet FG, Kindt R, Legendre P, Michin PR, O’Hara RB, Simpson GL, Solymos P, Henry M, Stevens H, Szoecs E, Wagner H (2012) Vegan: community ecology package [WWW document]. URL
  36. Parks KE, Mulligan M (2010) On the relationship between a resource based measure of geodiversity and broad scale biodiversity patterns. Biodivers Conserv 19:2751–2766CrossRefGoogle Scholar
  37. Peters DPC, Bestelmeyer BT, Turner MG (2007) Cross-scale interactions and changing pattern-process relationships: consequences for system dynamics. Ecosystems 10:790–796CrossRefGoogle Scholar
  38. Peters DPC, Gosz JR, Pockman WT, Small EE, Parmenter RR, Collins SL, Muldavin E (2006) Integrating patch and boundary dynamics to understand and predict biotic transitions at multiple scales. Landscape Ecol 21:19–33CrossRefGoogle Scholar
  39. R Core Team (2018) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. URL
  40. Read J, Jaffré T, Ferris JM, McCoy S, Hope GS (2006) Does soil determine the boundaries of monodominant rain forest with adjacent mixed rain forest and maquis on ultramafic soils in New Caledonia? J Biogeogr 33:1055–1065CrossRefGoogle Scholar
  41. Rietkerk M, Ketner P, Burger J, Hoorens B, Olff H (2000) Multiscale soil and vegetation patchiness along a gradient of herbivore impact in a semi-arid grazing system in West Africa. Plant Ecol 148:207–224CrossRefGoogle Scholar
  42. Robertson GP (2000) Geostatistics for Environmental Sciences: GS+ User’s Guide, Version 5. Gamma Design Software, MIGoogle Scholar
  43. Roem WJ, Berendse F (2000) Soil acidity and nutrient supply ratio as possible factors determining changes in plant species diversity in grassland and heathland communities. Biol Cons 92:151–161CrossRefGoogle Scholar
  44. Sankey TT, Montagne C, Graumlich L, Lawrence R, Nielsen J (2006) Lower forest-grassland ecotones and 20th Century livestock herbivory effects in northern Mongolia. Forest Ecol Manag 233:36–44CrossRefGoogle Scholar
  45. Shi D, Wang D (2005) Effects of various salt-alkaline mixed stresses on Aneurolepidium chinense (Trin.) Kitag. Plant Soil 271:15–26CrossRefGoogle Scholar
  46. Staal A, Flore BM (2015) Sharp ecotones spark sharp ideas: comment on “Structural, physiognomic and above-ground biomass variation in savanna-forest transition zones on three continents: how different are co-occurring savanna and forest formations?” by Veenendaal et al. (2015). Biogeosciences 12:5563–5566CrossRefGoogle Scholar
  47. Stein RA, Ludwig JA (1979) Vegetation and soil patterns on a Chihuahuan Desert bajada. Am Midl Nat 101:28–37CrossRefGoogle Scholar
  48. Stine MB, Resler LM, Campbell JB (2011) Ecotone characteristics of a southern Appalachian Mountain wetland. CATENA 86:57–65CrossRefGoogle Scholar
  49. Walkley A, Black IA (1934) An examination of Degtjareff method for determining soil organic matter and a proposed modification of the chromic acid titration method. Soil Sci 37:29–37CrossRefGoogle Scholar
  50. Wang D, Ba L (2008) Ecology of meadow steppe in northeast China. Rangel J 30:247–254CrossRefGoogle Scholar
  51. Wang D, Du J, Zhang B, Ba L, Hodgkinson KC (2017) Grazing intensity and phenotypic plasticity in the clonal grass Leymus chinensis. Rangel Ecol Manag 70:740–747CrossRefGoogle Scholar
  52. Wang L, Liu C, Alves DG, Frank DA, Wang D (2015) Plant diversity is associated with the amount and spatial structure of soil heterogeneity in meadow steppe of China. Landscape Ecol 30:1713–1721CrossRefGoogle Scholar
  53. Wang Z, Song K, Zhang B, Liu D, Ren C, Luo L, Yang T, Huang N, Hu L, Yang H, Liu Z (2009) Shrinkage and fragmentation of grasslands in the west Songnen Plain, China. Agric Ecosyst Environ 129:315–324CrossRefGoogle Scholar
  54. Wu J, David JL (2002) A spatially explicit hierarchical approach to modeling complex ecological systems: theory and applications. Ecol Model 153:7–26CrossRefGoogle Scholar
  55. Wu J, Levin SA (1994) A spatial patch dynamic modeling approach to pattern and process in an annual grassland. Ecol Monogr 64:447–464CrossRefGoogle Scholar
  56. Wu J, Loucks OL (1995) From balance of nature to hierarchical patch dynamics: a paradigm shift in ecology. Quart Rev Biol 70:439–466CrossRefGoogle Scholar
  57. Yang L, Han M, Zhou G, Li J (2007) The changes in water-use efficiency and stoma density of Leymus chinensis along Northeast China Transect. Acta Ecol Sin 27:16–23CrossRefGoogle Scholar
  58. Yang C, Shi D, Wang D (2008) Comparative effects of salt and alkali stresses on growth, osmotic adjustment and ionic balance of an alkali-resistant halophyte Suaeda glauca (Bge.). Plant Growth Regul 56:179–190CrossRefGoogle Scholar
  59. Zhang J, Li X, Liu Z, Li X, Gao Z, Mu C (2014) Growth forms of Leymus chinesis (Poaceae) at the different developmental stages of the natural population. Plant Species Biol 29:263–271CrossRefGoogle Scholar
  60. Zhong Z, Wang D, Zhu H, Wang L, Feng C, Wang Z (2014) Positive interactions between large herbivores and grasshoppers, and their consequences for grassland plant diversity. Ecology 95:1055–1064CrossRefGoogle Scholar

Copyright information

© Springer Nature B.V. 2019

Authors and Affiliations

  1. 1.State Environmental Protection Key Laboratory of Wetland Ecology and Vegetation Restoration / Key Laboratory of Vegetation Ecology, Ministry of Education, School of EnvironmentNortheast Normal UniversityChangchunChina
  2. 2.Department of Biological SciencesUniversity of BergenBergenNorway

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