Abstract
Vegetation patterns are a ubiquitous feature of water-deprived ecosystems. Despite the competition for the same limiting resource, coexistence of several plant species is commonly observed. We propose a two-species reaction–diffusion model based on the single-species Klausmeier model, to analytically investigate the existence of states in which both species coexist. Ecologically, the study finds that coexistence is supported if there is a small difference in the plant species’ average fitness, measured by the ratio of a species’ capabilities to convert water into new biomass to its mortality rate. Mathematically, coexistence is not a stable solution of the system, but both spatially uniform and patterned coexistence states occur as metastable states. In this context, a metastable solution in which both species coexist corresponds to a long transient (exceeding \(10^3\) years in dimensional parameters) to a stable one-species state. This behaviour is characterised by the small size of a positive eigenvalue which has the same order of magnitude as the average fitness difference between the two species. Two mechanisms causing the occurrence of metastable solutions are established: a spatially uniform unstable equilibrium and a stable one-species pattern which is unstable to the introduction of a competitor. We further discuss effects of asymmetric interspecific competition (e.g. shading) on the metastability property.
Similar content being viewed by others
References
Alfaro M, Izuhara H, Mimura M (2018) On a nonlocal system for vegetation in drylands. J Math Biol 77:1761–1793
Bastiaansen R, Jaïbi O, Deblauwe V, Eppinga MB, Siteur K, Siero E, Mermoz S, Bouvet A, Doelman A, Rietkerk M (2018) Multistability of model and real dryland ecosystems through spatial self-organization. In: Proceedings of the National Academy of Sciences, pp 201804771
Bates P, Xun J (1994) Metastable patterns for the Cahn–Hilliard equation, part I. J Differ Equ 111:421–457
Bates P, Xun J (1995) Metastable patterns for the Cahn–Hilliard equation: part II. Layer dynamics and slow invariant manifold. J Differ Equ 117:165–216
Baudena M, Rietkerk M (2013) Complexity and coexistence in a simple spatial model for arid savanna ecosystems. Theor Ecol 6:131–141
Baudena M, Boni G, Ferraris L, von Hardenberg J, Provenzale A (2007) Vegetation response to rainfall intermittency in drylands: results from a simple ecohydrological box model. Adv Water Resour 30:1320–1328
Baudena M, D’Andrea F, Provenzale A (2010) An idealized model for tree–grass coexistence in savannas: the role of life stage structure and fire disturbances. J Ecol 98:74–80
Bennett JJ, Sherratt JA (2018) Long-distance seed dispersal affects the resilience of banded vegetation patterns in semi-deserts. J Theor Biol. https://doi.org/10.1016/j.jtbi.2018.10.002
Borgogno F, D’Odorico P, Laio F, Ridolfi L (2009) Mathematical models of vegetation pattern formation in ecohydrology. Rev Geophys 47:RG1005
Buis E, Veldkamp A, Boeken B, van Breemen N (2009) Controls on plant functional surface cover types along a precipitation gradient in the Negev Desert of Israel. J Arid Environ 73:82–90
Callegaro C, Ursino N (2018) Connectivity of niches of adaptation affects vegetation structure and density in self-organized (dis-connected) vegetation patterns. Land Degrad Dev 29:2589–2594
Consolo G, Currò C, Valenti G (2019) Supercritical and subcritical Turing pattern formation in a hyperbolic vegetation model for flat arid environments. Phys D Nonlinear Phenom. https://doi.org/10.1016/j.physd.2019.03.006
Cornet A, Delhoume J, Montaña C (1988) Diversity and pattern in plant communities. In: During H, Werger M, Willems H (eds) Dynamics of striped vegetation patterns and water balance in the Chihuahuan Desert. SPB Academic Publishing, The Hague, pp 221–231
Corrado R, Cherubini AM, Pennetta C (2014) Early warning signals of desertification transitions in semiarid ecosystems. Phys Rev E Stat Nonlinear Soft Matter Phys 90:062705
Dakos V, Kéfi S, Rietkerk M, van Nes EH, Scheffer M (2011) Slowing down in spatially patterned ecosystems at the brink of collapse. Am Nat 177:E153–E166
Deblauwe V, Barbier N, Couteron P, Lejeune O, Bogaert J (2008) The global biogeography of semi-arid periodic vegetation patterns. Glob Ecol Biogeogr 17:715–723
Deblauwe V, Couteron P, Bogaert J, Barbier N (2012) Determinants and dynamics of banded vegetation pattern migration in arid climates. Ecol Monogr 82:3–21
d’Herbès J-M, Valentin C, Tongway DJ, Leprun J-C (2001) Banded vegetation patterns and related structures. In: Tongway DJ, Valentin C, Seghieri J (eds) Banded vegetation patterning in arid and semiarid environments: ecological processes and consequences for management. Springer, New York, pp 1–19
Dickovick JT (2014) Africa 2014–2015. World today (Stryker). Rowman & Littlefield Publishers, Lanham, p 374
D’Onofrio D, Baudena M, D’Andrea F, Rietkerk M, Provenzale A (2015) Treegrass competition for soil water in arid and semiarid savannas: the role of rainfall intermittency. Water Resour Res 51:169–181
Dunkerley D, Brown K (2002) Oblique vegetation banding in the Australian arid zone: implications for theories of pattern evolution and maintenance. J Arid Environ 51:163–181
Eigentler L, Sherratt JA (2018) Analysis of a model for banded vegetation patterns in semi-arid environments with nonlocal dispersal. J Math Biol 77:739–763
Eldridge D, Zaady E, Shachak M (2000) Infiltration through three contrasting biological soil crusts in patterned landscapes in the Negev, Israel. CATENA 40:323–336
Gandhi P, Werner L, Iams S, Gowda K, Silber M (2018) A topographic mechanism for arcing of dryland vegetation bands. J R Soc Interface 15:20180508
Gilad E, von Hardenberg J, Provenzale A, Shachak M, Meron E (2004) Ecosystem engineers: from pattern formation to habitat creation. Phys Rev Lett 93:098105
Gilad E, von Hardenberg J, Provenzale A, Shachak M, Meron E (2007a) A mathematical model of plants as ecosystem engineers. J Theor Biol 244:680–691
Gilad E, Shachak M, Meron E (2007b) Dynamics and spatial organization of plant communities in water-limited systems. Theor Popul Biol 72:214–230
Gowda K, Iams S, Silber M (2018) Signatures of human impact on selforganized vegetation in the Horn of Africa. Engl Sci Rep 8:1–8
Gowda K, Chen Y, Iams S, Silber M (2016) Assessing the robustness of spatial pattern sequences in a dryland vegetation model. Proc R Soc Lond A 472:20150893
Guttal V, Jayaprakash C (2007) Self-organization and productivity in semiarid ecosystems: implications of seasonality in rainfall. J Theor Biol 248:490–500
Hemming CF (1965) Vegetation arcs in Somaliland. J Ecol 53:57–67
HilleRisLambers R, Rietkerk M, van den Bosch F, Prins HHT, de Kroon H (2001) Vegetation pattern formation in semi-arid grazing systems. Ecology 82:50–61
Iron D, Ward MJ (2004) The stability and dynamics of hot-spot solutions to two one-dimensional microwave heating models. Anal Appl 02:21–70
Kealy BJ, Wollkind DJ (2012) A nonlinear stability analysis of vegetative turing pattern formation for an interaction–diffusion plant-surface water model system in an arid flat environment. Bull Math Biol 74:803–833
Kéfi S, Rietkerk M, Alados CL, Pueyo Y, Papanastasis V, ElAich A, de Ruiter P (2007) Spatial vegetation patterns and imminent desertification in Mediterranean arid ecosystems. Nature 449:213–217
Klausmeier CA (1999) Regular and irregular patterns in semiarid vegetation. Science 284:1826–1828
Kletter A, von Hardenberg J, Meron E, Provenzale A (2009) Patterned vegetation and rainfall intermittency. J Theor Biol 256:574–583
Kyriazopoulos P, Nathan J, Meron E (2014) Species coexistence by front pinning. Ecol Complex 20:271–281
Marasco A, Iuorio A, Carteni F, Bonanomi G, Tartakovsky DM, Mazzoleni S, Giannino F (2014) Vegetation pattern formation due to interactions between water availability and toxicity in plant–soil feedback. Bull Math Biol 76:2866–2883
Meron E (2012) Pattern-formation approach to modelling spatially extended ecosystems. Ecol Model 234:70–82
Meron E (2016) Pattern formation—a missing link in the study of ecosystem response to environmental changes. Math Biosci 271:1–18
Meron E (2018) From patterns to function in living systems: dryland ecosystems as a case study. Annu Rev Condens Matter Phys 9:79–103
Montaña C (1992) The colonization of bare areas in two-phase mosaics of an arid ecosystem. J Ecol 80:315–327
Montaña C, Lopez-Portillo J, Mauchamp A (1990) The response of two woody species to the conditions created by a shifting ecotone in an arid ecosystem. J Ecol 78:789–798
Moreno-de las Heras MM, Saco PM, Willgoose GR, Tongway DJ (2012) Variations in hydrological connectivity of Australian semiarid landscapes indicate abrupt changes in rainfall-use efficiency of vegetation. J Geophys Res G Biogeosci 117:G03009
Müller J (2013) Floristic and structural pattern and current distribution of tiger bush vegetation in Burkina Faso (West Africa), assessed by means of belt transects and spatial analysis. Appl Ecol Environ Res 11:153–171
Nathan J, von Hardenberg J, Meron E (2013) Spatial instabilities untie the exclusion-principle constraint on species coexistence. J Theor Biol 335:198–204
Pelletier JD, De Long SB, Orem CA, Becerra P, Compton K, Gressett K, Lyons-Baral J, McGuire LA, Molaro JL, Spinler JC (2012) How do vegetation bands form in dry lands? Insights from numerical modeling and field studies in southern Nevada, USA. J Geophys Res F Earth Surf 117:F04026
Penny GG, Daniels KE, Thompson SE (2013) Local properties of patterned vegetation: quantifying endogenous and exogenous effects. Philos Trans Soc R London Ser A 371:20120359
Potapov AB, Hillen T (2005) Metastability in chemotaxis models. J Dyn Differ Equ 17:293–330
Pueyo Y, Kéfi S, Alados CL, Rietkerk M (2008) Dispersal strategies and spatial organization of vegetation in arid ecosystems. Oikos 117:1522–1532
Pueyo Y, Kéfi S, Díaz-Sierra R, Alados C, Rietkerk M (2010) The role of reproductive plant traits and biotic interactions in the dynamics of semiarid plant communities. Theor Popul Biol 78:289–297
Reynolds JF, Smith DMS, Lambin EF, Turner BL, Mortimore M, Batterbury SPJ, Downing TE, Dowlatabadi H, Fernandez RJ, Herrick JE, Huber- Sannwald E, Jiang H, Leemans R, Lynam T, Maestre FT, Ayarza M, Walker B (2007) Global desertification: building a science for dryland development. Science 316:847–851
Rietkerk M, van de Koppel J (2008) Regular pattern formation in real ecosystems. Trends Ecol Evol 23:169–175
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 148:207–224
Rietkerk M, Boerlijst MC, van Langevelde F, HilleRisLambers R, van de Koppel J, Kumar L, Prins HHT, de Roos AM (2002) Self-organization of vegetation in arid ecosystems. Am Nat 160:524–530
Rietkerk M, Dekker SC, de Ruiter PC, van de Koppel J (2004) Self-organized patchiness and catastrophic shifts in ecosystems. Science 305:1926–1929
Rodriguez-Iturbe I, Porporato A, Ridolfi L, Isham V, Coxi DR (1999) Probabilistic modelling of water balance at a point: the role of climate, soil and vegetation. Proc R Soc Lond A 455:3789–3805
Saco PM, Moreno-de las Heras M, Keesstra S, Baartman J, Yetemen O, Rodriguez JF (2018) Vegetation and soil degradation in drylands: non linear feedbacks and early warning signals. Curr Opin Environ Sci Health 5:67–72
Salvucci GD (2001) Estimating the moisture dependence of root zone water loss using conditionally averaged precipitation. Water Resour Res 37:1357–1365
Scheiter S, Higgins S, Weissing AEFJ, Geber EMA (2007) Partitioning of root and shoot competition and the stability of savannas. Am Nat 170:587–601
Seghieri J, Galle S, Rajot J, Ehrmann M (1997) Relationships between soil moisture and growth of herbaceous plants in a natural vegetation mosaic in Niger. J Arid Environ 36:87–102
Serra-Diaz JM, Maxwell C, Lucash MS, Scheller RM, Laflower DM, Miller AD, Tepley AJ, Epstein HE, Anderson-Teixeira KJ, Thompson JR (2018) Disequilibrium of fire-prone forests sets the stage for a rapid decline in conifer dominance during the twenty-first century. Sci Rep 8:6749
Sheffer E, Hardenberg J, Yizhaq H, Shachak M, Meron E, Blasius B (2013) Emerged or imposed: a theory on the role of physical templates and selforganisation for vegetation patchiness. Ecol Lett 16:127–139
Sherratt JA (2005) An analysis of vegetation stripe formation in semi-arid landscapes. J Math Biol 51:183–197
Sherratt JA (2010) Pattern solutions of the Klausmeier model for banded vegetation in semi-arid environments I. Nonlinearity 23:2657–2675
Sherratt JA (2011) Pattern solutions of the Klausmeier model for banded vegetation in semi-arid environments II: patterns with the largest possible propagation speeds. Proc R Soc Lond A 467:3272–3294
Sherratt JA (2013a) History-dependent patterns of whole ecosystems. Ecol Complex 14:8–20
Sherratt JA (2013b) Pattern solutions of the Klausmeier model for banded vegetation in semi-arid environments III: the transition between homoclinic solutions. Physica D 242:30–41
Sherratt JA (2013c) Pattern solutions of the Klausmeier model for banded vegetation in semiarid environments IV: slowly moving patterns and their stability. SIAM J Appl Math 73:330–350
Sherratt JA (2013d) Pattern solutions of the Klausmeier model for banded vegetation in semiarid environments V: the transition from patterns to desert. SIAM J Appl Math 73:1347–1367
Sherratt JA, Lord GJ (2007) Nonlinear dynamics and pattern bifurcations in a model for vegetation stripes in semi-arid environments. Theor Popul Biol 71:1–11
Siero E (2018) Nonlocal grazing in patterned ecosystems. J Theor Biol 436:64–71
Siero E, Siteur K, Doelman A, van de Koppel J, Rietkerk M, Eppinga MB (2019) Grazing away the resilience of patterned ecosystems. Am Nat 193:472–480
Siteur K, Siero E, Eppinga MB, Rademacher JD, Doelman A, Rietkerk M (2014a) Beyond turing: the response of patterned ecosystems to environmental change. Ecol Complex 20:81–96
Siteur K, Eppinga MB, Karssenberg D, Baudena M, Bierkens MF, Rietkerk M (2014b) How will increases in rainfall intensity affect semiarid ecosystems? Water Resour Res 50:5980–6001
Sprugel DG (1991) Disturbance, equilibrium, and environmental variability: What is ‘natural’ vegetation in a changing environment? Biol Conserv 58:1–18
Svenning J-C, Sandel B (2013) Disequilibrium vegetation dynamics under future climate change. Am J Bot 100:1266–1286
Synodinos AD, Tietjen B, Jeltsch F (2015) Facilitation in drylands: modeling a neglected driver of savanna dynamics. Ecol Model 304:11–21
Thiery JM, D’Herbès J-M, Valentin C (1995) A model simulating the genesis of banded vegetation patterns in Niger. J Ecol 83:497–507
Thompson SE, Harman CJ, Heine P, Katul GG (2010) Vegetation-infiltration relationships across climatic and soil type gradients. J Geophys Res G Biogeosci 115:G02023
Tilman D (1982) Resource competition and community structure. Princeton University Press, Princeton, p 296
Tongway DJ, Ludwig JA (1990) Vegetation and soil patterning in semi-arid mulga lands of Eastern Australia. Aust J Ecol 15:23–34
Tzuk O, Ujjwal SR, Fernandez-Oto C, Seifan M, Meron E (2019) Interplay between exogenous and endogenous factors in seasonal vegetation oscillations. Sci Rep 9:354
United Nations Convention to Combat Desertification (2017) The global land outlook. Version first edition, Bonn, Germany
United Nations Food and Agriculture Organization (2005) Livestock sector briefs
Ursino N, Callegaro C (2016) Diversity without complementarity threatens vegetation patterns in arid lands. Ecohydrology 9:1187–1195
Ursino N, Contarini S (2006) Stability of banded vegetation patterns under seasonal rainfall and limited soil moisture storage capacity. Adv Water Resour 29:1556–1564
Valentin C, d’Herbès J, Poesen J (1999) Soil and water components of banded vegetation patterns. CATENA 37:1–24
van der Stelt S, Doelman A, Hek G, Rademacher JDM (2013) Rise and fall of periodic patterns for a generalized Klausmeier–Gray–Scott model. J Nonlinear Sci 23:39–95
White LP (1971) Vegetation stripes on sheet wash surfaces. J Ecol 59:615–622
Worrall GA (1959) The Butana grass patterns. J Soil Sci 10:34–53
Zelnik YR, Kinast S, Yizhaq H, Bel G, Meron E (2013) Regime shifts in models of dryland vegetation. Philos Trans R Soc Lond Ser A 371:20120358
Zelnik YR, Gandhi P, Knobloch E, Meron E (2018) Implications of tristability in pattern-forming ecosystems. Chaos Interdiscip J Nonlinear Sci 28:033609
Zimmerman JK, Comita LS, Thompson J, Uriarte M, Brokaw N (2010) Patch dynamics and community metastability of a subtropical forest: compound effects of natural disturbance and human land use. Landsc Ecol 25:1099–1111
Author information
Authors and Affiliations
Corresponding author
Additional information
Publisher's Note
Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Lukas Eigentler was supported by The Maxwell Institute Graduate School in Analysis and its Applications, a Centre for Doctoral Training funded by the UK Engineering and Physical Sciences Research Council (Grant EP/L016508/01), the Scottish Funding Council, Heriot-Watt University and the University of Edinburgh.
Rights and permissions
About this article
Cite this article
Eigentler, L., Sherratt, J.A. Metastability as a Coexistence Mechanism in a Model for Dryland Vegetation Patterns. Bull Math Biol 81, 2290–2322 (2019). https://doi.org/10.1007/s11538-019-00606-z
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11538-019-00606-z