The study of local and regional mechanisms driving spatial patterns in beta diversity is essential to the understanding of biodiversity. We aimed to predict the roles of multiple mechanisms operating at regional to fine spatial scales in structuring the beta diversity in a mountainous semi-arid rangeland of the Khorassan-Kopet Dagh floristic province located in NE Iran. We evaluated the relative contributions of three main filters, at the regional (stochastic but spatially structured dispersal filter), local (abiotic filter), and fine (biotic filter) spatial scales on beta diversity across communities. We partitioned beta diversity constrained by spatial, environmental, and biotic variables in 23 communities studied here, and used simple RDA and RDA-based variation partitioning to assess the contributions of studied filters on beta diversity. Moreover, spatial autocorrelation analyses were used to test neutral theory predictions. The relative contributions of the studied ecological filters explained 55% of variation in beta diversity. Although differences in the explained variations between unique fractions are low, the abiotic and biotic filters (signifying the niche-based processes) represent stronger effects directly and indirectly (via impact on significantly of other ecological filters) than dispersal (signifying neutral processes) on beta diversity. In addition, 45% of the variation in beta diversity was not explained by the studied ecological filters. In conclusion, independent and shared impacts of processes at different spatial scales determine beta diversity in our plant communities. However, unexplained variation in beta diversity requires further study of other facets of biodiversity and community assembly processes.
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Alejandro GFB, Chinchilla FA, Magrach A, Romero V, Reyos M, Velilla M, Amador-Vargas Serrano JM, S, (2009) Slope orientation enhances the nurse effect of a paramo shrub, Hypericum irazuense (Hypericaceae) in Costa Rica. J Trop Ecol 25:331–335
Anderson MJ, Crist TO, Chase JM, Vellend M, &, et al (2011) Navigating the multiple meanings of beta diversity: a roadmap for the practicing ecologist. Ecol Lett 14:19–28
Armas C, Pugnaire FI (2011) Belowground zone of influence in a tussock grass species. Acta Oecol 37:284–289
BA Carlsson, Callaghan TV (1991) Positive plant interactions in tundra vegetation and the importance of shelter. J Ecol 79:973–983. https://doi.org/10.2307/2261092
Badano EI, Cavieres LA (2006) Ecosystem engineering across ecosystems: do engineer species sharing common features have generalized or idiosyncratic effects on species diversity? J Biogeogr 33:304–313
Bertness MD, Callaway R (1994) Positive interactions in communities. Trends in Ecol Evol 9(5):191–193. https://doi.org/10.1016/0169-5347(94)90088-4
Borcard D, Legendre P (2002) All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrices. Ecol Model 153:51–68
Borcard D, Legendre P, Drapeau P (1992) Partialling out the spatial component of ecological variation. Ecology 73:1045–1055
Bowker MA, Maestre FT, Escolar C (2010) Biological crusts as a model system for examining the biodiversity–ecosystem function relationship in soils. Soil Biol Biochem 42:405–417
Bremner JM (1996) In: Sparks DL, et al. (eds) Nitrogen-Total Methods of soil analysis. Science Society of America Inc, American Society of Agronomy Inc, Madison, pp 1085–1122
Brooker RW, Maestre FT, Callaway RM, Lortie CL, Cavieres LA, Kunstler G, Liancourt P, Tielbo¨ rger K, Travis JMJ, Anthelme F, Armas C, Coll L, Corcket E, Delzon S, Forey E, Kikvidze Z, Olofsson J, Pugnaire FI, Saccone P, Schiffer K, Seifan M, Touzard B, Michalet R, (2008) Facilitation in plant communities: the past, the present, and the future. J Ecol 96:18–34
Butterfield BJ, Cavieres LA, Callaway RM, Cook BJ, Kikvidze Z, Lortie CJ, Brooker RW (2013) Alpine cushion plants inhibit loss of phylogenetic diversity in severe environments. Ecol Lett 16:478–486. https://doi.org/10.1111/ele.12070
Callaway RM (2007) Positive interactions and interdependence in plant communities. Springer, New York
Callaway RM, Kikodze D, Kikvidze Z (2000) Facilitation by unpalatable weeds may conserve plant diversity in overgrazed meadows in the Caucasus Mountains. Oikos 89:275–282
Callaway RM, Brooker RW, Choler P, Kikvidze Z, Lortie CJ, Michalet R et al (2002) Positive interactions among alpine plants increase with stress. Nature 417:844–848. https://doi.org/10.1038/nature00812
Carr SC, Robertson KM, Platt WJ, Peet RK (2009) A model of geographical, environmental and regional variation in vegetation composition of pyrogenic grasslands of Florida. J Biogeogr 36:1600–1612
Cavender-Bares J, Kozak K, Fine P, Kembel S (2009) The merging of community ecology and phylogenetic biology. Ecol Lett 12:693–715
Chamberlain SA, Bronstein JL, Rudgers JA (2014) How context dependent are species interactions? Ecol Lett 17(7):881–890. https://doi.org/10.1111/ele.12279
Chang LW, Zelen D, Li CF, Chiu ST, Hsieh C-F (2013) Better environmental data may reverse conclusions about niche- and dispersal-based processes in community assembly. Ecology 94:2145–2151
Chao A, Chazdon RL, Colwell RK, Shen T (2005) A new statistical approach for assessing similarity of species composition with incidence and abundance data. Ecol Lett 8:148–159
Chase JM (2007) Drought mediates the importance of stochastic community assembly. PNAS 104:17430–17434
Choler P, Michalet R, Callaway RM (2001) Facilitation and competition on gradients in alpine plant communities. Ecology 82:3295–3308
Cortina J, Maestre FT (2005) Plant effects on soils in drylands: implications for community dynamics and ecosystem restoration. In: Binkley D, Menyailo O (eds) Tree species effects on soils: implications for global change. NATO Science Series, Kluwer Academic Publishers, Dordrecht, 85–118.
Diniz-Filho JAF, Siqueira T, Padial AA, Rangel TF, Landeiro VL, Bini LM (2012) Spatial autocorrelation analysis allows disentangling the balance between neutral and niche processes in meta communities. Oikos 121(2):201–210
Dray S (2010) SpacemakeR: Spatial modelling. R package version 0.0–5. https://r-forge.r-project.org/sedar. Accessed 2 June 2013
Dray S, Legendre P, Peres-Neto PR (2006) Spatial modelling: a comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecol Model 196:483–493
Erktan A, Rey F (2013) Linking sediment trapping efficiency with morphological traits of Salix tiller barriers on marly gully floors under ecological rehabilitation. Ecol Eng 51:212–220
Farzam M, Ejtehadi H (2016) Effects of drought and canopy facilitation on plant diversity and abundance in a semiarid mountainous rangeland. J Plant Ecol 10(4):626–633
Gotelli NJ (2000) Null model analysis of species co-occurrence patterns. Ecol 81:2606–2621
Gotelli NJ, Graves GR (1996) Null models in Ecology. Smithsonian Institution Press, Washington DC
Harrison S, Ross SJ, Lawton JH (1992) Beta diversity on geographic gradients in Britain. J Anim Ecol 61:151–158
He Q, Bertness MD, Altieri AH (2013) Global shifts towards positive species interactions with increasing environmental stress. Ecol Lett 16(5):695–706
Hijmans RJ, Cameron SE, Parra JL et al (2005) Very high resolution interpolated climate surfaces for global land areas. Int J Clim 25:1965–1978
Jankju M, Delavari A, Ganjali A (2008) Interseeding Bromus kopetdaghensis, in shrublands. Rangeland J Iran Soc Range Manag 2:314–328
Kraft NJB, Ackerly DD (2014) Assembly of plant communities. In: Monson RK (ed) Ecology and the environment, the plant sciences, vol 8. Springer, New York, NY, pp 67–88
Kraft NJB, Adler PB, Godoy O, James EC, Fuller S, Levine JM (2015) Community assembly, coexistence and the environmental filtering metaphor. Funct Ecol 29:592–599
Laliberté E, Paquette A, Legendre P, Bouchard A (2009) Assessing the scale-specific importance of niches and other spatial processes on beta diversity: a case study from a temperate forest. Oecologia 159(2):377–388
Legendre P (1993) Spatial autocorrelation: trouble or new paradigm. Ecology 74:1659–1673
Legendre P, De Caceres M (2013) Beta diversity as the variance of community data: dissimilarity coefficients and partitioning. Ecol Lett 16:951–963
Legendre P, Gallagher ED (2001) Ecologically meaningful transformations for ordination of species data. Oecologia 129:271–280
Legendre P, Legendre L (1998) Numerical Ecology, 2nd edn. Elsevier, Amsterdam
Legendre P, Legendre L (2012) Numerical ecology, Third English edition. Elsevier, Amsterdam
Legendre P, Borcard D, Peres-Neto PR (2005) Analyzing beta diversity: partitioning the spatial variation of community composition data. Ecol Monogr 75:435–450
Legendre P, Borcard D, Roberts DW (2012) Variation partitioning involving orthogonal spatial eigenfunction submodels. Ecol 93(5):1234–1240
Lennon JJ, Koleff P, Greenwood JJD, Gaston KJ (2001) The geographical structure of British bird distributions: diversity, spatial turnover and scale. J Anim Ecol 70:966–979
Lessard JP, Belmaker J, Myers JA, Chase JM, Rahbek C (2012) Inferring local ecological processes amid species pool influences. Trends Ecol Evol. 27(11):600–607
Li X, Sun H (2017) Phylogenetic pattern of alpine plants along latitude and longitude in Hengduan Mountains Region. Plant Divers 39:37–43
Lin G, Stralberg D, Gong G, Huang Z, Ye W, Wu L (2013) Separating the effects of environment and space on tree species distribution: from population to community. PLoS ONE 8(2):e56171. https://doi.org/10.1371/journal.pone.0056171
Lopez RP, Valdivia S, Rivera ML, Rios RS (2013) Co-occurrence patterns along a regional aridity gradient of the subtropical andes do not support stress gradient hypotheses. PLoS ONE 8:1–10
Lopez-Angulo J, Swenson NG, Cavieres LA (2018) Interactions between abiotic gradients determine functional and phylogenetic diversity patterns in Mediterranean type climate mountains in the Andes. J Veg Sci 29(2):245–254
Lortie CJ, Brooker RW, Choler P, Kikvidze Z, Michalet R, Callaway FI, Pugnaire RM (2004) Rethinking plant community theory. Oikos 107:433–438
Luiz AM, Leao-Pires TA, Sawaya RJ (2016) Geomorphology drives amphibian beta diversity in Atlantic forest Lowlands of southeastern Brazil. PLoS ONE 11(5):e0153977
Luzuriaga AL, Sanchez AN, Maestre FT, Escudero A (2012) Assemblage of a semi-arid annual plant community: abiotic and biotic filters act hierarchically. PLoS ONE 7(7):e41270
Maestre FT, Bautista S, Cortina J (2003) Positive, negative and net effects in grass–shrub interactions in Mediterranean semiarid grasslands. Ecology 84:3186–3197
MAPA (1994) Métodos Oficiales de Análisis. Tomo III. Servicio de Publicaciones del Ministerio de Agricultura. Pescay Alimentación, Madrid, pp. 662. (In French)
Mayfield MM, Levine JM (2010) Opposing effects of competitive exclusion on the phylogenetic structure of communities. Eco lett 13:1085–1093
McCune B, Grace J (2002) Multivariate analysis of ecological communities. MjM Software, Gleneden Beach
Memariani F, Akhani H, Joharchi MR (2016) Endemic plants of Khorassan-Kopet Dagh floristic province in Irano-Turanian region: diversity, distribution patterns and conservation status. Phytotaxa 249(1):031–117
Memariani F et al (2016) A review of plant diversity, vegetation and phytogeography of the Khorassan-Kopet Dagh floristic province in the Irano-Turanian region (northeastern Iran – southern Turkmenistan). Phytotaxa 249(1):8–30
Michale R (2006) Highlighting the multiple drivers of change in interactions along stress gradients. New Phytol 173:3–6
Miranda JD, Armas C, Padilla FM, Pugnaire FI (2011) Climatic change and rainfall patterns: effects on semi-arid plant communities of the Iberian Southeast. J Arid Environ 75:1302–1309
Mori AS et al (2015) Null model approaches to evaluating the relative role of different assembly processes in shaping ecological communities. Oecologia 178:261–273
Moura MR, Villalobos F, Costa GC, Garcia PCA (2016) Disentangling the role of climate, topography and vegetation in species richness gradients. PLoS ONE 11(3):e0152468
Muster C, Meyer M, Sattler T (2014) Spatial arrangement overrules environmental factors to structure native and non-native assemblages of synanthropic harvestmen. PLoS ONE 9(3):e90474
Myers JA et al (2013) Beta-diversity in temperate and tropical forests reflects dissimilar mechanisms of community assembly. Ecol Lett 16:151–157
Økland RH (2003) Partitioning the variation in a plot-byspecies data matrix that is related to n sets of explanatory variables. J Veg Sci 14:693–700
Økland RH, Eilersten O (1994) Canonical correspondence analysis with variation partitioning: some comments and an application. J Veg Sci 5:117–126
Oksanen J, Blanchet FG, Kindt R, Legendre P, Minchin PR, O’Hara RB, et al. (2015) Vegan: Community Ecology Package. R package version 2.0–9. https://CRAN.R-project.org/package=vegan. Accessed 16 Sept 2015
Pashirzad M, Ejtehadi H, Vaezi J, Sheferson RP (2018) Spatial scale-dependent phylogenetic signal in species distributions along geographic and elevation gradients in a mountainous rangeland. Ecol Evol 8(21):10364–10373. https://doi.org/10.1002/ece3.4293
Pashirzad M, Ejtehadi H, Vaezi J, Shefferson RP (2019) Plant-plant interactions influence phylogenetic diversity at multiple spatial scales in a semi-arid mountain rangeland. Oecologia 189:1–11
Pausas JG, Verdú M, (2010) The jungle of methods for evaluating phenotypic and phylogenetic structure of communities. Bioscience 60:614–625
Peres-Neto PR, Legendre P (2010) Estimating and controlling for spatial structure in the study of ecological communities. Glob Ecol Biogeogr 19(2):174–184
Peres-Neto PR, Legendre P, Dray S, Borcard D (2006) Variation partitioning of species data matrices: estimation and comparison of fractions. Ecology 87:2614–2625
Peters H, O’Leary BC, Hawkins JP, Carpenter KE, Roberts CM (2013) Conus: first comprehensive conservation red list assessment of a marine gastropod mollusc genus. PLoS ONE 8(12):e83353. https://doi.org/10.1371/journal.pone.0083353
Pirani A, Rabeler RK (2017) Nomenclatural notes on Acanthophyllum (Caryophylleae, Caryophyllaceae). Phytotaxa 303(2):197–198
Piston N, Schob C, Armas C, Prieto I, Pugnaire F (2016) Contribution of co-occurring shrub species to community richness and phylogenetic diversity along an environmental gradient. Perspect Plant Ecol Evol Syst 19:30–39
Pottier J, Dubuis A, Pellissier L, Maiorano L, Rossier L, Randin CF, Guisan A (2013) The accuracy of plant assemblage prediction from species distribution models varies along environmental gradients. Glob Ecol Biogeogr 22(1):52–63
Qian H, Chen SH, Zhang JL (2017) Disentangling environmental and spatial effects on phylogenetic structure of angiosperm tree communities in China. Sci Rep 7:5864
Qiao X, Li Q, Jiang Q, Lu J, Franklin S et al (2015) Beta diversity determinants in Badagongshan, a subtropical forest in central China. Sci Rep 5:17043
Rajaniemi TK, Turkington R, Goldberg D (2009) Community-level consequences of species interactions in an annual plant community. J Veg Sci 20:836–846
Ricklefs RE (2008) Disintegration of the ecological community. Am Nat 172(6):741–750
Shipley B (2015) Describing, explaining and predicting community assembly: a convincing trait-based case study. J Veg Sci 26:615–616
Soininen J, Lennon JJ, Hillebrand H (2007) Multivariate analysis of beta diversity across organisms and environments. Ecology 88(11):2830–2838
Soliveres S, Eldridge DJ, Maestre FT, Bowker MA, Tighe M, Escudero A (2011) Microhabitat amelioration and reduced competition among understory plants as drivers of facilitation across environmental gradients: towards a unifying framework. Perspect Plant Ecol Evol Syst 13:247–258
Soliveres S, Torices R, Maestre F (2012) Environmental conditions and biotic interactions acting together promote phylogenetic randomness in semiarid plant communities: new methods help to avoid misleading conclusions. J Veg Sci 23:822–836
Stein A, Gerstner K, Kreft H (2014) Environmental heterogeneity as a universal driver of species richness across taxa, biomes and spatial scales. Ecol Lett 17(7):866–880
Tello JS et al (2015) Elevational gradients in β-diversity reflect variation in the strength of local community assembly mechanisms across spatial scales. PLoS ONE 10:e0121458
Tonkin JD, Altermatt F, Finn DS, Heino J, Olden JD, Pauls SU (2018) The role of dispersal in river network metacommunities: patterns, processes, and pathways. Freshw Biol 63(1):141–163. https://doi.org/10.1111/fwb.13037
Valiente-Banuet A, Vital A, Verdú M, Callaway RM, (2006) Modern quaternary plant lineages promote diversity through facilitation of ancient Tertiary lineages. Proc Nat Acad Sci USA 103:16812–16817
Vavrek MJ (2015) Fossil package.
Vellend M (2010) Conceptual synthesis in community ecology. Q Rev Biol 85(2):183–206
Verdú M, Rey PJ, Alcantara JM, Siles G, Valiente-Banuet A (2009) Phylogenetic signatures of facilitation and competition in successional communities. J Ecol 97:1171–1180
Walkley A, Black IA (1934) An examination of the Degtjareff method for determining soil organic matter, and a proposed modification of the chromic acid titration method. Soil Sci 37:29–38
Wang B et al (2015) Relationship between topography and the distribution of understory vegetation in a Pinus massoniana forest in Southern China. Int Soil Water Conserv Res 3:291–304
Weinstein BG, Catherine H, Parra JL (2017) The role of environment, dispersal and competition in explaining reduced co-occurrence among related species. PLoS ONE 12(11):e0185493. https://doi.org/10.1371/journal.pone.0185493
Yang J, Swenson NJ, Zhang G et al (2015) Local-scale partitioning of functional and phylogenetic beta diversity in a tropical tree assemblage. Sci Rep 5:12731
The authors wish to thank the Ferdowsi University of Mashhad for funding this research under the Grant No.3/41572.
This study was funded by Ferdowsi University of Mashhad (Grant Number 3/41572).
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Communicated by Lauchlan Fraser.
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Pashirzad, M., Ejtehadi, H., Vaezi, J. et al. Multiple processes at different spatial scales determine beta diversity patterns in a mountainous semi-arid rangeland of Khorassan-Kopet Dagh floristic province, NE Iran. Plant Ecol 220, 829–844 (2019). https://doi.org/10.1007/s11258-019-00957-6
- Multiple processes
- Spatial scales
- Beta diversity patterns
- Biotic filter
- Abiotic filter
- Dispersal filter