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Oecologiastronger in tropical than temperate forests because ofhigher numbers of specialist predators and pathogens in theformer (Janzen 1970; Connell 1971; Coley and Barone1996; Givnish 1999; Harms et al. 2000; Dyer et al. 2007),and the latitudinal gradient in tree diversity has beenhypothesized to be caused by decreasing Janzen–Connelleffects with increasing latitude. However, studies havefound that density-dependent mortality is as common intemperate forests as in tropical forests. For example, HilleRis Lambers et al. (2002) found the proportion of speciesaffected by negative density dependence at the seed andseedling stages in a North American temperate forest wasequivalent to that reported for tropical forests, though theyacknowledged that the magnitude of density-dependenteffects may be higher in tropical forests. Our study ofdensity dependence in Liangshui FDP also supports theidea that density dependence may be as important intemperate forests as in tropical forests, and densitydependence alone is unlikely to explain latitudinal treediversity differences in the world’s forests (but see Johnsonet al. 2012).ConclusionIn combination with previous studies from tropical andsubtropical forests, our results from an old-growth temperateforest indicate that significant negative effects oflocal conspecific density occur at multiple life stages intree communities. This suggests that density dependenceplays an important role in enhancing community diversityof forests in different latitudes. Although we found negativeconspecific effects at multiple stages, species exhibitingnegative density dependence at one life stage did notnecessarily exhibit it at other stages. Thus, analyses thatfocus on a single life stage may underestimate the prevalenceand importance of density dependence in tree communities.Similarly, our results suggest that studies that failto take into account confounding factors, such as habitatheterogeneity and species-level variation, may also mischaracterizethe role of density dependence in shapingplant communities. Therefore, we recommend that futurestudies take habitat heterogeneity and other potentiallyconfounding factors into account and also test for effects ofconspecific neighbors across all life stages.Acknowledgments This study was financially supported by theNational Natural Science Foundation of China (No. 30770350,31270473), and the Natural Science Foundation of HeilongjiangProvince, China (No. ZJN0706). Many critical comments on themanuscript were received in the 2011 Smithsonian CTFS/SIGEO andCForBio Workshop in Changbaishan (US NSF grant DEB-1046113).We are grateful to Keping Ma and Zhanqing Hao for their efforts inorganizing that workshop, and the technological support fromSmithsonian Tropical Research Institute. We also thank ThorstenWiegand, Stephan Getzin and Yan Zhu for providing useful suggestions.The conduction of this study complied with the current laws ofChina.Open Access This article is distributed under the terms of theCreative Commons Attribution License which permits any use, distribution,and reproduction in any medium, provided the originalauthor(s) and the source are credited.ReferencesBarot S, Gignoux J, Menaut JC (1999) Demography of a savannapalm tree: predictions from comprehensive spatial patternsanalysis. Ecology 80:1987–2000Bates D, Maechler M, Dai B (2008) lme4: linear mixed-effectsmodels using S4 classes. 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