Conspecific negative density dependence (CNDD) could be one of the most important local-scale mechanisms shaping plant species coexistence. However, the spatial and temporal changes in the strength CNDD and the implications for the plant diversity remain unknown. We used 10 years of seedling data, in a seasonal tropical rainforest, to discover how CNDD influences tree seedling survival across habitats and seasons. We also evaluated the relation between CNDD and species diversity. We found the strength of CNDD in the valley habitat was significantly stronger than in ridge habitat in rainy season, but not significantly different in dry season. Corresponding to expectations of CNDD as mechanism of diversity maintenance, seedling species diversity was significantly higher in valley habitat than in ridge habitat and significantly correlated with CNDD. Additionally, conspecific and heterospecific seedling neighbour densities positively affected the survival of tree seedlings, but heterospecific adult neighbour density had a weak effect. Our study finds that CNDD varied significantly across habitats and was correlated with local seedling diversity. Our results highlight the importance of CNDD in driving species diversity at the local scale. Recognizing the spatial and temporal variation in the strength of CNDD will aid efforts to model and understand species coexistence.
This is a preview of subscription content, access via your institution.
Buy single article
Instant access to the full article PDF.
Tax calculation will be finalised during checkout.
Subscribe to journal
Immediate online access to all issues from 2019. Subscription will auto renew annually.
Tax calculation will be finalised during checkout.
Aide TM (1992) Dry season leaf production—an escape from herbivory. Biotropica 24:532–537. https://doi.org/10.2307/2389016
Bagchi R, Gallery RE, Gripenberg S, Gurr SJ, Narayan L, Addis CE, Freckleton RP, Lewis OT (2014) Pathogens and insect herbivores drive rainforest plant diversity and composition. Nature 506:85–88. https://doi.org/10.1038/nature12911
Bates D, Maechler M, Bolker B, Walker S (2014) lme4: linear mixed-effects models using Eigen and S4. R package version 1
Bell T, Freckleton RP, Lewis OT (2006) Plant pathogens drive density-dependent seedling mortality in a tropical tree. Ecol Lett 9:569–574. https://doi.org/10.1111/j.1461-0248.2006.00905.x
Bever JD, Platt TG, Morton ER (2012) Microbial population and community dynamics on plant roots and their feedbacks on plant communities. Annu Rev Microbiol 66:265–283. https://doi.org/10.1146/annurev-micro-092611-150107
Brenes-Arguedas T, Coley PD, Kursar TA (2009) Pests vs. drought as determinants of plant distribution along a tropical rainfall gradient. Ecology 90:1751–1761. https://doi.org/10.1890/08-1271.1
Bunker DE, Carson WP (2005) Drought stress and tropical forest woody seedlings: effect on community structure and composition. J Ecol 93:794–806. https://doi.org/10.1111/j.1365-2745.2005.01019.x
Cao M, Zou XM, Warren M, Zhu H (2006) Tropical forests of Xishuangbanna, China. Biotropica 38:306–309. https://doi.org/10.1111/j.1744-7429.2006.00146.x
Carson WP, Anderson JT, Leigh EG, Schnitzer SA (2008) Challenges associated with testing and falsifying the Janzen-Connell hypothesis: a review and critique. In: Carson WP, Schnitzer SA (eds) Tropical forest community ecology. Blackwell Publishing Ltd, Oxford, pp 210–241
Chen L, Mi X, Comita LS, Zhang L, Ren H, Ma K (2010) Community-level consequences of density dependence and habitat association in a subtropical broad-leaved forest. Ecol Lett 13:695–704. https://doi.org/10.1111/j.1461-0248.2010.01468.x
Chesson P (1994) Multispecies competition in variable environments. Theor Popul Biol 45:227–276
Chesson P (2000) Mechanisms of maintenance of species diversity. Annu Rev Ecol Syst 31:343–366. https://doi.org/10.1146/annurev.ecolsys.31.1.343
Coley PD (1998) Possible effects of climate change on plant/herbivore interactions in moist tropical forests. Clim Change 39:455–472
Coley PD, Barone JA (1996) Herbivory and plant defenses in tropical forests. Annu Rev Ecol Syst 27:305–335. https://doi.org/10.1146/annurev.ecolsys.27.1.305
Comita LS, Hubbell SP (2009) Local neighborhood and species' shade tolerance influence survival in a diverse seedling bank. Ecology 90:328–334. https://doi.org/10.1890/08-0451.1
Comita LS, Muller-Landau HC, Aguilar S, Hubbell SP (2010) Asymmetric density dependence shapes species abundances in a tropical tree community. Science 329:330–332. https://doi.org/10.1126/science.1190772
Comita LS, Queenborough SA, Murphy SJ, Eck JL, Xu K, Krishnadas M, Beckman N, Zhu Y (2014) Testing predictions of the Janzen-Connell hypothesis: a meta-analysis of experimental evidence for distance- and density-dependent seed and seedling survival. J Ecol 102:845–856. https://doi.org/10.1111/1365-2745.12232
Condit R (1998) Tropical forest census plots: methods and results from Barro Colorado Island, Panama and a comparison with other plots. Springer Science & Business Media, New York
Connell J (1971) On the role of natural enemies in preventing competitive exclusion in some marine animals and in rain forest trees, Dynamics of Population. In: Den Boer PJ, Gradwell GR (eds) Dynamics of populations centre for agricultural publishing and documentation. Wageningen, The Netherlands, pp 298–312
Crawford KM, Bauer JT, Comita LS, Eppinga MB, Johnson DJ, Mangan SA, Queenborough SA, Strand AE, Suding KN, Umbanhowar J, Bever JD (2019) When and where plant-soil feedback may promote plant coexistence: a meta-analysis. Ecol Lett 22:1274–1284. https://doi.org/10.1111/ele.13278
Detto M, Visser MD, Wright SJ, Pacala SW (2019) Bias in the detection of negative density dependence in plant communities. Ecol Lett. https://doi.org/10.1111/ele.13372
Dirzo R, Boege K (2008) Patterns of herbivory and defense in tropical dry and rain forests. In: Carson WP, Schnitzer SA (eds) Tropical forest community ecology. Blackwell Publishing Ltd, Oxford, pp 63–78
Forrister DL, Endara MJ, Younkin GC, Coley PD, Kursar TA (2019) Herbivores as drivers of negative density dependence in tropical forest saplings. Science 363:1213–1216. https://doi.org/10.1126/science.aau9460
Gilbert GS, Parker IM (2016) The evolutionary ecology of plant disease: a phylogenetic perspective. Annu Rev Phytopathol 54:549–578. https://doi.org/10.1146/annurev-phyto-102313-045959
Harms KE, Wright SJ, Calderon O, Hernandez A, Herre EA (2000) Pervasive density-dependent recruitment enhances seedling diversity in a tropical forest. Nature 404:493–495. https://doi.org/10.1038/35006630
He FL, Duncan RP (2000) Density-dependent effects on tree survival in an old-growth Douglas fir forest. J Ecol 88:676–688. https://doi.org/10.1046/j.1365-2745.2000.00482.x
HilleRisLambers J, Adler P, Harpole W, Levine J, Mayfield M (2012) Rethinking community assembly through the lens of coexistence theory. Annu Rev Ecol Evol Syst 43:227–248. https://doi.org/10.1146/annurev-ecolsys-110411-160411
Hossain M, Veneklaas EJ, Hardy GES, Poot P (2019) Tree host-pathogen interactions as influenced by drought timing: linking physiological performance, biochemical defence and disease severity. Tree Physiol 39:6–18. https://doi.org/10.1093/treephys/tpy113
Hu YH (2010) A study on the habitat variation and distribution patterns of tree species in a 20-ha forest dynamics plot at Bubeng, Xishuangbanna, SW China. PhD thesis, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences
Janzen DH (1970) Herbivores and the number of tree species in tropical forests. Am Nat 104:501–528. https://doi.org/10.1086/282687
Johnson DJ, Beaulieu WT, Bever JD, Clay K (2012) Conspecific negative density dependence and forest diversity. Science 336:904–907. https://doi.org/10.1126/science.1220269
Johnson DJ, Bourg NA, Howe R, McShea WJ, Wolf A, Clay K (2014) Conspecific negative density-dependent mortality and the structure of temperate forests. Ecology 95:2493–2503
Johnson DJ, Condit R, Hubbell SP, Comita LS (2017) Abiotic niche partitioning and negative density dependence drive tree seedling survival in a tropical forest. Proc R Soc B 284:8. https://doi.org/10.1098/rspb.2017.2210
LaManna JA, Walton ML, Turner BL, Myers JA (2016) Negative density dependence is stronger in resource-rich environments and diversifies communities when stronger for common but not rare species. Ecol Lett 19:657–667. https://doi.org/10.1111/ele.12603
LaManna JA, Mangan SA, Alonso A, Bourg NA, Brockelman WY, Bunyavejchewin S, Chang LW, Chiang JM, Chuyong GB, Clay K, Condit R, Cordell S, Davies SJ, Furniss TJ, Giardina CP, Gunatilleke IAUN, Gunatilleke CVS, He FL, Howe RW, Hubbell SP, Hsieh CF, Inman-Narahari FM, Janik D, Johnson DJ, Kenfack D, Korte L, Kral K, Larson AJ, Lutz JA, McMahon SM, McShea WJ, Memiaghe HR, Nathalang A, Novotny V, Ong PS, Orwig DA, Ostertag R, Parker GG, Phillips RP, Sack L, Sun IF, Tello JS, Thomas DW, Turner BL, Diaz DMV, Vrska T, Weiblen GD, Wolf A, Yap S, Myers JA (2017) Plant diversity increases with the strength of negative density dependence at the global scale. Science 356:1389–1392
Leigh EG, Davidar P, Dick CW, Puyravaud JP, Terborgh J, ter Steege H, Wright SJ (2004) Why do some tropical forests have so many species of trees? Biotropica 36:447–473. https://doi.org/10.1111/j.1744-7429.2004.tb00342.x
Liang M, Liu X, Gilbert GS, Zheng Y, Luo S, Huang F, Yu S (2016) Adult trees cause density-dependent mortality in conspecific seedlings by regulating the frequency of pathogenic soil fungi. Ecol Lett 19:1448–1456. https://doi.org/10.1111/ele.12694
Lin L, Comita LS, Zheng Z, Cao M (2012) Seasonal differentiation in density-dependent seedling survival in a tropical rain forest. J Ecol 100:905–914. https://doi.org/10.1111/j.1365-2745.2012.01964.x
Lu J, Johnson DJ, Qiao X, Lu Z, Wang Q, Jiang M (2015) Density dependence and habitat preference shape seedling survival in a subtropical forest in central China. J Plant Ecol 8:568–577
Martinez-Camilo R, Gonzalez-Espinosa M, Ramirez-Marcial N, Cayuela L, Angel Perez-Farrera M (2018) Tropical tree species diversity in a mountain system in southern Mexico: local and regional patterns and determinant factors. Biotropica 50:499–509. https://doi.org/10.1111/btp.12535
Martinkova Z, Honek A (2011) Asymmetrical intraspecific competition in Echinochloa crus-galli is related to differences in the timing of seedling emergence and seedling vigour. Plant Ecol 212:1831–1839. https://doi.org/10.1007/s11258-011-9953-8
Meinzer FC, Andrade JL, Goldstein G, Holbrook NM, Cavelier J, Wright SJ (1999) Partitioning of soil water among canopy trees in a seasonally dry tropical forest. Oecologia 121:293–301. https://doi.org/10.1007/s004420050931
Murphy SJ, Wiegand T, Comita LS (2017) Distance-dependent seedling mortality and long-term spacing dynamics in a neotropical forest community. Ecol Lett 20:1469–1478. https://doi.org/10.1111/ele.12856
O'Brien MJ, Reynolds G, Ong R, Hector A (2017) Resistance of tropical seedlings to drought is mediated by neighbourhood diversity. Nat Ecol Evol 1:1643–1648. https://doi.org/10.1038/s41559-017-0326-0
Paine CET, Harms KE, Schnitzer SA, Carson WP (2008) Weak competition among tropical tree seedlings: Implications for species coexistence. Biotropica 40:432–440. https://doi.org/10.1111/j.1744-7429.2007.00390.x
Peters HA (2003) Neighbour-regulated mortality: the influence of positive and negative density dependence on tree populations in species-rich tropical forests. Ecol Lett 6:757–765. https://doi.org/10.1046/j.1461-0248.2003.00492.x
Pu X, Zhu Y, Jin G (2017) Effects of local biotic neighbors and habitat heterogeneity on seedling survival in a spruce-fir valley forest, northeastern China. Ecol Evol 7:4582–4591. https://doi.org/10.1002/ece3.3030
Queenborough SA, Burslem D, Garwood NC, Valencia R (2007) Neighborhood and community interactions determine the spatial pattern of tropical tree seedling survival. Ecology 88:2248–2258. https://doi.org/10.1890/06-0737.1
R Core Team (2018) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna
Song QH, Tan ZH, Zhang YP, Cao M, Sha LQ, Tang Y, Liang NS, Schaefer D, Zhao JF, Zhao JB, Zhang X, Yu L, Deng XB (2013) Spatial heterogeneity of soil respiration in a seasonal rainforest with complex terrain. Iforest-Biogeosci For 6:65–72. https://doi.org/10.3832/ifor0681-006
Song X, Hogan JA, Lin L, Wen H, Cao M, Yang J (2018a) Canopy openness and topographic habitat drive tree seedling recruitment after snow damage in an old-growth subtropical forest. For Ecol Manage 429:493–502. https://doi.org/10.1016/j.foreco.2018.07.038
Song X, Johnson DJ, Cao M, Umana MN, Deng XB, Yang XF, Zhang WF, Yang J (2018b) The strength of density-dependent mortality is contingent on climate and seedling size. J Veg Sci 29:662–670. https://doi.org/10.1111/jvs.12645
Spear ER, Coley PD, Kursar TA (2015) Do pathogens limit the distributions of tropical trees across a rainfall gradient? J Ecol 103:165–174. https://doi.org/10.1111/1365-2745.12339
Sterck F, Markesteijn L, Schieving F, Poorter L (2011) Functional traits determine trade-offs and niches in a tropical forest community. P Natl Acad Sci USA 108:20627–20632. https://doi.org/10.1073/pnas.1106950108
Svenning JC, Fabbro T, Wright SJ (2008) Seedling interactions in a tropical forest in Panama. Oecologia 155:143–150. https://doi.org/10.1007/s00442-007-0884-y
Swinfield T, Lewis OT, Bagchi R, Freckleton RP (2012) Consequences of changing rainfall for fungal pathogen-induced mortality in tropical tree seedlings. Ecol Evol 2:1408–1413. https://doi.org/10.1002/ece3.252
Terborgh J (2012) Enemies maintain hyperdiverse tropical forests. Am Nat 179:303–314. https://doi.org/10.1086/664183
Wright JS (2002) Plant diversity in tropical forests: a review of mechanisms of species coexistence. Oecologia 130:1–14
Wu J, Swenson NG, Brown C, Zhang C, Yang J, Ci X, Li J, Sha L, Cao M, Lin L (2016) How does habitat filtering affect the detection of conspecific and phylogenetic density dependence? Ecology 97:1182–1193. https://doi.org/10.1890/14-2465.1
Zhang C, Yang J, Sha L, Ci X, Li J, Cao M, Brown C, Swenson NG, Lin L (2017) Lack of phylogenetic signals within environmental niches of tropical tree species across life stages. Sci Rep. https://doi.org/10.1038/srep42007
Zhu Y, Comita LS, Hubbell SP, Ma K (2015) Conspecific and phylogenetic density-dependent survival differs across life stages in a tropical forest. J Ecol 103:957–966. https://doi.org/10.1111/1365-2745.12414
Zhu Y, Queenborough SA, Condit R, Hubbell SP, Ma KP, Comita LS (2018) Density-dependent survival varies with species life-history strategy in a tropical forest. Ecol Lett 21:506–515. https://doi.org/10.1111/ele.12915
This research was supported by the National Natural Science Foundation of China (31800353, 31870410 and 31670442), the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB31000000), the CAS 135 program (no. 2017XTBG-T01), and the West Light Foundation of Chinese Academy of Sciences. We thank Xishuangbanna Station for Tropical Rain Forest Ecosystem Studies of Xishuangbanna Tropical Botanical Garden for assistance with the field work.
Conflict of interest
The authors declare that they have no conflict of interest.
Communicated by Marie-Josée Fortin.
Electronic supplementary material
Below is the link to the electronic supplementary material.
About this article
Cite this article
Song, X., Zhang, W., Johnson, D.J. et al. Conspecific negative density dependence in rainy season enhanced seedling diversity across habitats in a tropical forest. Oecologia 193, 949–957 (2020). https://doi.org/10.1007/s00442-020-04729-2
- Janzen–Connell hypothesis
- Plant–soil feedback
- Plant–plant interaction
- Species diversity