The small-scale variation of herb-layer community structure in a riparian mixed forest

  • Y. O. Zhukova Oles Honchar Dnipro National University
  • N. V. Yorkina Bogdan Khmelnitsky Melitopol State Pedagogical University
  • V. S. Budakova Bogdan Khmelnitsky Melitopol State Pedagogical University
  • O. M. Kunakh Oles Honchar Dnipro National University
Keywords: overstorey structure; soil properties; spatial pattern; phytoindication; scalogram; spatial eigenvector mapping.


The ground vegetation layer is the most diverse plant community in forest ecosystems. We have shown the role of spatial variables, soil properties and overstorey structure in spatial variation of the herb-layer community in a riparian mixed forest . The research was conducted in the "Dnipro-Orils’kiy" Nature Reserve (Ukraine). The research polygon was located in the forest in the floodplain of the River Protich, which is a left tributary of the River Dnipro. Plant abundance was quantified by measuring cover within an experimental polygon. The experimental polygon consisted of 7 transects, each comprising 15 test points. The distance between the rows in the site was 3 m. At the site we established a plot of 45 × 21 m, with 105 subplots of 3 × 3 m organized in a regular grid. A list of vascular plant species was composed for each 3 × 3 m subplot along with visual estimates of species cover projection. The plant community was represented by 43 species, of which 18.6% were phanerophytes, 39.5% were hemicryptophytes, 9.3% were therophytes, 7.0% were geophytes. An overall test of random labelling revealed the total nonrandom distribution of the tree stems within the site. The species-specific test of random labelling showed the nonrandom segregated distribution of Acer tataricum, Pyrus communis, Quercus robur, and Ulmus laevis. Crataegus monogyna and Euonymus europaeus were distributed randomly. The nearest neighbour of Acer tataricum was less likely to be Ulmus laevis. There was no direct spatial connection between Acer tataricum and other trees. Crataegus monogyna, Pyrus communis, Quercus robur and Euonymus europaeus were not segregated from all other species. The nearest neighbour of Ulmus laevis was less likely to be Acer tataricum. Constrained correspondence analysis (CCA) was applied as ordination approach. The forward selection procedure allowed us to select 6 soil variables which explain 28.3% of the herb-layer community variability. The list of the important soil variables includes soil mechanical impedance (at the depth 0–5, 30–35, 75–80, and 95–10 cm), soil moisture, and soil bulk density. The variation explained by pure spatial variables accounted for 11.0 %. The majority of the tree-distance structured variation in plant community composition was broad-scaled. The spatial scalograms were left-skewed asymmetric. Significant relationship was found between the pure spatial component of the community variation and a number of phytoindicator estimations, most important of which were the variability of damping and humidity. Tree stand was obseerved to be a considerable factor structuring both the herb-layer community and spatial variation of the physical properties of soil.


Aiba, M., Takafumi, H., & Hiura, T. (2012). Interspecific differences in determinants of plant species distribution and the relationships with functional traits. Journal of Ecology, 100(4), 950–957.

Andivia, E., Fernández, M., Alejano, R., & Vázquez-Piqué, J. (2015). Tree patch distribution drives spatial heterogeneity of soil traits in cork oak woodlands. Annals of Forest Science, 72(5), 549–559.

Angers, D. A., & Caron, J. (1998). Plant-induced changes in soil structure: Processes and feedbacks. Biogeochemistry, 42(1/2), 55–72.

Baddeley, A., & Turner, R. (2005). Spatstat : An R package for analyzing spatial point patterns. Journal of Statistical Software, 12(6), 1–42.

Barbier, S., Gosselin, F., & Balandier, P. (2008). Influence of tree species on understory vegetation diversity and mechanisms involved-A critical review for temperate and boreal forests. Forest Ecology and Management, 254(1), 1–15.

Binkley, D., & Giardina, C. (1998). Why do tree species affect soils? The warp and woof of tree-soil interactions. In: Plant-induced soil changes: Processes and feedbacks. Springer Netherlands, Dordrecht. Pp. 89–106.

Blanchet, F. G., Legendre, P., & Borcard, D. (2008). Forward selection of explanatory variables. Ecology, 89(9), 2623–2632.

Blank, L., & Carmel, Y. (2012). Woody vegetation patch types affect herbaceous species richness and composition in a Mediterranean ecosystem. Community Ecology, 13(1), 72–81.

Borcard, D., & Legendre, P. (2002). All-scale spatial analysis of ecological data by means of principal coordinates of neighbour matrices. Ecological Modelling, 153(1–2), 51–68.

Borsukevish, L. V., & Onishenko, V. A. (2018). Moist and occasionally flooded oak-elm forests. In: Kuzemko, A. A. (Ed.). National habitat catalogue of Ukraine. FOP Y. I. Klymenko, Kyiv. Pp. 262–263.

Bratton, S. P. (1976). Resource division in an understory herb community: Responses to temporal and microtopographic gradients. The American Naturalist, 110(974), 679–693.

Breshears, D. D., Rich, P. M., Barnes, F. J., & Campbell, K. (1997). Overstory-imposed heterogeneity in solar radiation and soil moisture in a semiarid woodland. Ecological Applications, 7(4), 1201–1215.

Brygadyrenko, V. (2015). Evaluation of the ecological niche of some abundant species of the subfamily Platyninae (Coleoptera, Carabidae) against the background of eight ecological factors. Folia Oecologica, 42, 1–18.

Brygadyrenko, V. (2019). Evaluation of ecological niches of abundant species of Poecilus and Pterostichus (Coleoptera: Carabidae) in forests of steppe zone of Ukraine. Entomologica Fennica, 27(2), 81–100.

Brygadyrenko, V. V. (2016). Effect of canopy density on litter invertebrate community structure in pine forests. Ekológia (Bratislava), 35(1), 90–102.

Burns, A. R., Stephens, W. Z., Stagaman, K., Wong, S., Rawls, J. F., Guillemin, K., & Bohannan, B. J. (2016). Contribution of neutral processes to the assembly of gut microbial communities in the zebrafish over host development. The ISME Journal, 10(3), 655–664.

Burrascano, S., Sabatini, F. M., & Blasi, C. (2011). Testing indicators of sustainable forest management on understorey composition and diversity in southern Italy through variation partitioning. Plant Ecology, 212(5), 829–841.

Canty, A., & Ripley, B. (2016). Boot: Bootstrap R (S-Plus) functions. R Package Version, 1, 3–18.

Chang, L. W., Zelený, D., Li, C. F., Chiu, S. T., & Hsieh, C. F. (2013). Better environmental data may reverse conclusions about niche- and dispersal-based processes in community assembly. Ecology, 94(10), 2145–2151.

Chase, J. M. (2014). Spatial scale resolves the niche versus neutral theory debate. Journal of Vegetation Science, 25(2), 319–322.

Chave, J. (2004). Neutral theory and community ecology. Ecology Letters, 7(3), 241–253.

Chudomelová, M., Zelený, D., & Li, C. F. (2017). Contrasting patterns of fine-scale herb layer species composition in temperate forests. Acta Oecologica, 80, 24–31.

Cottenie, K. (2005). Integrating environmental and spatial processes in ecological community dynamics. Ecology Letters, 8(11), 1175–1182.

Dallas, T., & Drake, J. M. (2014). Relative importance of environmental, geographic, and spatial variables on zooplankton metacommunities. Ecosphere, 5(9), 104.

Damgaard, C. F., & Irvine, K. M. (2019). Using the beta distribution to analyse plant cover data. Journal of Ecology, 107(6), 2747–2759.

De la Cruz, M. (2008). Metodos para analizar datos puntuales. In: Maestre, F. T., Escudero, A., & Bonet, A. (Eds.). Introduccion al analisis espacial de datos en ecologia y ciencias ambientales: Metodos y aplicaciones, asociacion espanola de ecologia terrestre. Universidad Rey Juan Carlos y Caja de Ahorros del Mediterraneo, Madrid. Pp. 76–127.

Didukh, Y. P. (2011). The ecological scales for the species of Ukrainian flora and their use in synphytoindication. Phytosociocentre, Kyiv.

Dixon, P. M. (2002). Nearest-neighbor contingency table analysis of spatial segregation for several species. Écoscience, 9(2), 142–151.

Dray, S., Pélissier, R., Couteron, P., Fortin, M. J., Legendre, P., Peres-Neto, P. R., Bellier, E., Bivand, R., Blanchet, F. G., De Cáceres, M., Dufour, A. B., Heegaard, E., Jombart, T., Munoz, F., Oksanen, J., Thioulouse, J., & Wagner, H. H. (2012). Community ecology in the age of multivariate multiscale spatial analysis. Ecological Monographs, 82(3), 257–275.

Faly, L. I., & Brygadyrenko, V. V. (2014). Patterns in the horizontal structure of litter invertebrate communities in windbreak plantations in the steppe zone of the Ukraine. Journal of Plant Protection Research, 54(4), 414–420.

Frelich, L. E., Machado, J.-L., & Reich, P. B. (2003). Fine-scale environmental variation and structure of understorey plant communities in two old-growth pine forests. Journal of Ecology, 91(2), 283–293.

Gazol, A., & Ibáñez, R. (2010). Plant species composition in a temperate forest: Multi-scale patterns and determinants. Acta Oecologica, 36(6), 634–644.

Gilliam, F. S. (2007). The ecological significance of the herbaceous layer in temperate forest ecosystems. BioScience, 57(10), 845–858.

Greacen, E. L., Farrell, D. A., & Cockroft, B. (1968). Soil resistance to metal probes and plant roots. Transactions of the 9th Congress of the International Society of Soil Science, 1, 769–779.

Gritsan, Y. I., Kunakh, O. M., Dubinina, J. J., Kotsun, V. I., & Tkalich, Y. I. (2019). The catena aspect of the landscape diversity of the “Dnipro-Orilsky” Natural Reserve. Journal of Geology, Geography and Geoecology, 28(3), 417–431.

Hubbell, S. (2001). The unified neutral theory of biodiversity and biogeography. Princeton University Press.

Hubbell, S. P. (2005). Neutral theory in community ecology and the hypothesis of functional equivalence. Functional Ecology, 19(1), 166–172.

Jones, C. G., Lawton, J. H., & Shachak, M. (1994). Organisms as ecosystem engineers. Oikos, 69(3), 373.

Karst, J., Gilbert, B., & Lechowicz, M. J. (2005). Fern community assembly: The roles of chance and the environment at local and intermediate scales. Ecology, 86(9), 2473–2486.

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., & Legendre, L. (2012). Numerical ecology. Ch. 6. Multidimensional qualitative data. Developments in Environmental Modelling, 24, 337–424.

Legendre, P., Mi, X., Ren, H., Ma, K., Yu, M., Sun, I. F., & He, F. (2009). Partitioning beta diversity in a subtropical broad-leaved forest of China. Ecology, 90(3), 663–674.

Lososová, Z., Šmarda, P., Chytrý, M., Purschke, O., Pyšek, P., Sádlo, J., Tichý, L., & Winter, M. (2015). Phylogenetic structure of plant species pools reflects habitat age on the geological time scale. Journal of Vegetation Science, 26(6), 1080–1089.

Lyon, J., & Sharpe, W. E. (2003). Impacts of hay‐scented fern on nutrition of northern red oak seedlings. Journal of Plant Nutrition, 26(3), 487–502.

Medvedev, V. V. (2009). Soil penetration resistance and penetrographs in studies of tillage technologies. Eurasian Soil Science, 42(3), 299–309.

Nettesheim, F. C., Garbin, M. L., Rajão, P. H. M., Araujo, D. S. D., & Grelle, C. E. V. (2018). Environment is more relevant than spatial structure as a driver of regional variation in tropical tree community richness and composition. Plant Ecology and Diversity, 11(1), 27–40.

Oijen, D., Feijen, M., Hommel, P., Ouden, J., & Waal, R. (2005). Effects of tree species composition on within‐forest distribution of understorey species. Applied Vegetation Science, 8(2), 155–166.

Oksanen, J., Blanchet, F. G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P. R., O’Hara, R. B., Simpson, G. L., Solymos, P., Stevens, M. H. H., Szoecs, E., & Wagner, H. (2019). Vegan: Community Ecology Package. R package version 2.5-6.

Oksanen, J., Blanchet, F. G., Kindt, R., Legendre, P., Minchin, P. R., O’Hara, R. B., Simpson, G. L., Solymos, P., Stevens, M. H. H., & Wagner, H. (2018). Community Ecology Package. R package version 2.5-2.

Paluch, J., & Gruba, P. (2012). Effect of local species composition on topsoil properties in mixed stands with silver fir (Abies alba Mill.). Forestry, 85(3), 413–426.

Pausas, J. G., & Austin, M. P. (2001). Patterns of plant species richness in relation to different environments: An appraisal. Journal of Vegetation Science, 12(2), 153–166.

R Core Team (2020). A language and environment for statistical computing. In: R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.

Rinklebe, J., & Langer, U. (2006). Microbial diversity in three floodplain soils at the Elbe River (Germany). Soil Biology and Biochemistry, 38(8), 2144–2151.

Rosindell, J., Hubbell, S. P., & Etienne, R. S. (2011). The unified neutral theory of biodiversity and biogeography at age ten. Trends in Ecology and Evolution, 26(7), 340–348.

Schindler, S., O’Neill, F. H., Biró, M., Damm, C., Gasso, V., Kanka, R., van der Sluis, T., Krug, A., Lauwaars, S. G., Sebesvari, Z., Pusch, M., Baranovsky, B., Ehlert, T., Neukirchen, B., Martin, J. R., Euller, K., Mauerhofer, V., & Wrbka, T. (2016). Multifunctional floodplain management and biodiversity effects: A knowledge synthesis for six European countries. Biodiversity and Conservation, 25(7), 1349–1382.

Schnitzler, A., Hale, B. W., & Alsum, E. (2005). Biodiversity of floodplain forests in Europe and Eastern North America: A comparative study of the Rhine and Mississippi Valleys. Biodiversity and Conservation, 14(1), 97–117.

Siefert, A., Ravenscroft, C., Althoff, D., Alvarez-Yépiz, J. C., Carter, B. E., Glennon, K. L., Heberling, J. M., Jo, I. S., Pontes, A., Sauer, A., Willis, A., & Fridley, J. D. (2012). Scale dependence of vegetation-environment relationships: A meta-analysis of multivariate data. Journal of Vegetation Science, 23(5), 942–951.

Stohlgren, T. J., Owen, A. J., & Lee, M. (2000). Monitoring shifts in plant diversity in response to climate change: A method for landscapes. Biodiversity and Conservation, 9(1), 65–86.

Talbot, C. J., Bennett, E. M., Cassell, K., Hanes, D. M., Minor, E. C., Paerl, H., Raymond, P. A., Vargas, R., Vidon, P. G., Wollheim, W., & Xenopoulos, M. A. (2018). The impact of flooding on aquatic ecosystem services. Biogeochemistry, 141(3), 439–461.

Ter Braak, C. J. F. (1986). Canonical correspondence analysis: A new eigenvector technique for multivariate direct gradient analysis. Ecology, 67(5), 1167–1179.

ter Braak, C. J. F., & Prentice, I. C. (1988). A theory of gradient analysis. Advances in Ecological Research, 18(C), 271–317.

Thieler, A. M., Fried, R., & Rathjens, J. (2016). RobPer: An R package to calculate periodograms for light curves based on robust regression. Journal of Statistical Software, 69, 9.

Tuomisto, H. (2003). Dispersal, environment, and floristic variation of western amazonian forests. Science, 299(5604), 241–244.

von Oheimb, G., & Härdtle, W. (2009). Selection harvest in temperate deciduous forests: Impact on herb layer richness and composition. Biodiversity and Conservation, 18(2), 271–287.

Wälder, K., Wälder, O., Rinklebe, J., & Menz, J. (2008). Estimation of soil properties with geostatistical methods in floodplains. Archives of Agronomy and Soil Science, 54(3), 275–295.

Ward, J., Tockner, K., Arscott, D., & Claret, C. (2002). Riverine landscape diversity. Freshwater Biology, 47(4), 517–539.

Weiher, E., Freund, D., Bunton, T., Stefanski, A., Lee, T., & Bentivenga, S. (2011). Advances, challenges and a developing synthesis of ecological community assembly theory. Philosophical Transactions of the Royal Society B: Biological Sciences, 366(1576), 2403–2413.

Yakovenko, V. M., Dubinina, J. J., & Zhukova, Y. O. (2019). Spatial heterogeneity of the physical properties of the soil in the floodplain of the small river. Agrology, 2(4), 219‒228.

Zhukov, A. V. (2015a). Phytoindicator estimation of the multidimensional scaling of the plant community structure. Biological Bulletin of Bogdan Chmelnitskiy Melitopol State Pedagogical University, 1(1), 69–93.

Zhukov, A. V. (2015b). Influence of usual and dual wheels on soil penetration resistance: The GIS-approach. Biological Bulletin of Bogdan Chmelnitskiy Melitopol State Pedagogical University, 5(3), 73–100.

Zhukov, A. V., & Zadorozhnaya, G. А. (2016). Spatio-temporal dynamics of the penetration resistance of recultivated soils formed after open cast mining. Visnyk of Dnipropetrovsk University, Biology, Ecology, 24(2), 324–331.

Zhukov, A. V., Andrusevich, K. V., Lapko, K. V., & Sirotina, V. O. (2015). Geostatistical estimation of soil aggregate structure as a composite variable. Biological Bulletin of Bogdan Chmelnitskiy Melitopol State Pedagogical University, 5(3), 101–121.

Zhukov, O. V., & Gubanova, N. L. (2015a). Diversity and dynamics of amphibians in floodplain ecosystems of the Samara river. Visnyk of Dnipropetrovsk University, Biology, Ecology, 23(1), 66–73.

Zhukov, O. V., & Gubanova, N. L. (2015b). Dynamic stability of communities of amphibians in short-term-floodedforest ecosystems. Visnyk of Dnipropetrovsk University, Biology, Ecology, 23(2), 161–171.

Zhukov, O. V., Kunah, O. M., Dubinina, Y. Y., & Ganzha, D. S. (2017). Diversity and phytoindication ability of plant community. Ukrainian Journal of Ecology, 7(4), 81–99.

Zhukov, O. V., Kunah, O. M., Dubinina, Y. Y., Fedushko, M. P., Kotsun, V. I., Zhukova, Y. O., & Potapenko, O. V. (2019). Tree canopy affects soil macrofauna spatial patterns on broad- and meso-scale levels in an Eastern European poplar-willow forest in the floodplain of the River Dnipro. Folia Oecologica, 46(2), 101–114.

Zhukov, O., Kunah, O., Dubinina, Y., Zhukova, Y., & Ganzha, D. (2019). The effect of soil on spatial variation of the herbaceous layer modulated by overstorey in an Eastern European poplar-willow forest. Ekologia Bratislava, 38(3), 253–272.

Zinke, P. J. (1962). The pattern of influence of individual forest trees on soil properties. Ecology, 43(1), 130–133.


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