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   Journal Publications


103 Chu, C.-J., Bartlett, M., Wang, Y.S., He, F., Weiner, J., Chave, J. and Sack, L. 2015. Does climate directly influence NPP globally? GLobal Change Biology (in press).


102 Mayor, S.J., Cahill, J., He, F. and Boutin, S. 2015. Scaling disturbance instead of richness to better understand anthropogenic impacts on biodiversity. PLoS ONE 10(5): e0125579 (doi: 10.1371/journal.pone.0125579).


101 Blanchet, G., Legendre, P., He, F. 2015. A new cost-effective approach to survey ecological communities. Oikos (in press).


100 He, F., Huang, S.M. and Zhang, J. 2015. An inconvenient truth about the long-term forest change in western Canada: competition, not climate, is the primary driving force; Reply to Price et al. Proceedings of the National Academy of Sciences ( (pdf).


99 Xu, H., Li, Y.-D., Liu, S.-R., Zang, R.G., He, F. and SpenceJ.R. 2015. Partial recovery of a tropical rainforest a half century after clear-cut and selective logging. Journal of Applied Ecology (in press).


98 Bin, Y., Spence, J., Wu, L.-F., Li, B.-H., Hao, Z.-Q., Ye, W.H. and He, F. 2015. Species-habitat associations and demographic rates of forest trees. Ecography (in press).


97 Zhang, J., Huang, S.M. and He, F. 2015. Half-century evidence from western Canada shows forest dynamics is primarily driven by competition followed by climate. Proceedings of the National Academy of Sciences 112: 4009-4014 (pdf).


96 Blanchet, F.G., Legendre, P., Bergeron, J.A.C. and He, F. 2014. Consensus RDA across dissimilarity coefficients for canonical ordination of community composition data. Ecological Monographs 84:491-511 (pdf).


95 Zhang, J., Huang, S., Hogg, E.H., Lieffers, V., Qin, Y. and He, F. 2014. Estimating spatial variation in Alberta forest biomass from a combination of forest inventory and remote sensing data. Biogeosciences 11:2793-2808.


94 Yin, D.-Y. and He, F. 2014. A simple method for estimating species abundance from occurrence maps. Methods in Ecology and Evolution 5:336-343..


93 Zhang, J., Mayor, S.J. and He, F. 2014. Does disturbance regime change community assembly of angiosperm plant communities in the boreal forest? Journal of Plant Ecology 7:188-201.



Gibson, L., Lynam, A.J., Bradshaw, C.J.A., He, F., Bickford, D., Woodruff, D.S., Bumrungsri, S. and Laurance, W.F. 2013. Near-complete extinction of native small mammal fauna 25 years after forest fragmentation. Science 341:1508-1510.


91 Shen, G.-C., Wiegand, T., Mi, X.-C. and He, F. Quantifying spatial phylogenetic structures of fully stem-mapped plant communities. Methods in Ecology and Evolution 4:1132–1141.



Cáceres, M.D., Legendre, P. and He, F. 2013. Dissimilarity measurements and the size structure of ecological communities. Methods in Ecology and Evolution 4:1167–1177.


89 Shen, G.-C., He, F., Waagepetersen, R., Sun, I-F., Hao, Z.-Q., Chen, Z.-S. and Yu, M.-J. 2013. Quantifying effects of habitat heterogeneity and other clustering processes on spatial distributions of tree species. Ecology 94:2436-2443.



Yan, E.-R., Wang, X.-H., Chang, S.X. and He, F. 2013. Scaling relationships among twig size, leaf size and leafing intensity in a successional series of subtropical forests. Tree Physiology 33:609-617.



He, F. and Hubbell, S.P. 2013. Estimating extinction from species-area relationships: Why the numbers do not add up. Ecology 94:1905-1912 (pdf).


Zhang, J., Kissling, D. and He, F. 2013. Local forest structure, climate and human disturbance determine regional distribution of boreal bird species richness in Alberta, Canada. Journal of Biogeography 40:1131-1142.


Guillaume, F.B., Bergeron, J.C.A., Spence, J.R. and He, F. 2013.Landscape effects of disturbance, habitat heterogeneity and spatial autocorrelation on a ground beetle (Carabidae) assemblage in mature boreal forest. Ecography 36:636-647.

  84 Wiegand, T. He, F. and Hubbell, S.P. 2013. A systematic comparison of summary characteristics for quantifying point patterns in ecology. Ecography 36:92-103.

Mayor, S.J., Cahill, J.C., He, F., Sólymos, P., Boutin, S. 2012. Regional boreal biodiversity peaks at intermediate human disturbance. Nature Communications 3:1142 (pages 1-6; pdf).

  82 Xu, H., Liu, S.-R, Li, Y.D., Zang, R.-G. and He, F. 2012. Assessing non-parametric and area-based methods for estimating regional species richness. Journal of Vegetation Science 23:1006-1012.

Cáceres, M.D., Legendre, P., Valencia, R., Cao, M., Chang, L.-W., Chuyong, G., Condit, R., Hao, Z.-Q., Hsieh, C.-F., Hubbell, S., Kenfack, D., Ma, K.-P., Mi, X.-C., Noor, Md. N.S., Kassim, A.R., Ren, H.-B., Su, S.-H., Sun, I-F., Thomas, D., Ye, W.-H. and He, F. 2012. The variation of tree beta diversity across a global network of forest plots. Global Ecology and Biogeography 12:1191-1202.


Ding, Y., Zang, R., Letcher, S.G., Liu, S. and He, F. 2012. Disturbance regime changes the trait distribution, phylogenetic structure, and community assembly of tropical rain forests. Oikos 121:1263-1270.


He, F. 2012. Area based assessment of extinction risk. Ecology 93: 974-980 (pdf).


Rosindell, J., Hubbell, S.P., He, F., Harmon, L.J. and Etienne, R.S. 2012. The case for ecological neutral theory. Trends in Ecology and Evolution 27: 203-208.


Ding, Y., Zang, R., Liu, S., He, F. and Letcher, S.G. 2012. Recovery of woody plant diversity in tropical rain forests in southern China after logging and shifting cultivation. Biological Conservation 145:225-233.


He, F. and Hubbell, S.P. 2012. Geometry and scale in species-area relationships; Extinction and climate change: a reply. Nature 482:E5-E6 (doi:10.1038/nature10859).


Yu, S.X., Zhang, D.Y. and He, F. 2012. The multitude of biodiversity: methods, theories and applications (Editorial). Journal of Plant Ecology 5:1-2.


He, F., Zhang, D.Y. and Lin, K. 2012. Coexistence of nearly neutral species. Journal of Plant Ecology 5:72-81.


Zhang, D.Y., Zhang, B.Y., Lin, K., Jiang, X.H., Tao, Y., Hubbell, S.P., He, F. and Ostling, A. 2012. Demographic trade-offs determine species abundance and diversity. Journal of Plant Ecology 5:82-88.


He, F. and Hubbell, S.P. 2011. Species–area relationships always overestimate extinction rates from habitat loss. Nature 473:368-371 (pdf; News&Views).


Hwang, W.-H. and He, F. 2011. Estimating abundance from presence-absence maps. Methods in Ecology and Evolution (DOI: 10.1111/j.2041-210X.2011.00105.x).


Zillio, T. and He, F. 2010. Modeling spatial aggregation of finite populations. Ecology 91:3698-3706 (pdf).


He, F. 2010. Maximum entropy, logistic regression, and species abundance. Oikos 119:578-582 (pdf).


Zillio, T. and He, F. 2010. Inferring species abundance distribution across spatial scales. Oikos 119:71-80 (pdf).


Gray, L. and He, F. 2009 Spatial point-pattern analysis for detecting density-dependent competition in a boreal chronosequence of Alberta. Forest Ecology and Management 259:98-106 (pdf).


Niu, K.C., Liu, Y.N., Shen, Z.H., He, F. and Fang, J.Y. 2009. Community assembly: the relative importance of neutral theory and niche theory. Biodiversity Science 17:579-593.


Yavitt, J.B., Harms, K.E., Garcia, M.N., Wright, S.J., He, F. and Mirabello, S.J. 2009. Spatial heterogeneity of soil chemical properties in a lowland tropical moist forest, Panama. Australian Journal of Soil Research 47:674-687.


Morlon, H., White, E.P., Etienne, R.S., Green, J.L., Ostling, A., Alonso, D., Enquist, B.J., He, F. , Hurlbert, A., Magurran, A.E., Maurer, B.A., McGill, B.J., Olff, H., Storch, D. and Zillio, T. 2009. Taking species abundance distributions beyond individuals. Ecology Letters 12:488-501.



Hu, X.-S., He, F. and Hubbell, S.P. 2009. Communtiy differentiation on landscapes: drift, migration, and speciation. Oikos 118:1515-1523 (pdf).



He, F. 2009. Price of prosperity: economic development and biological conservation in China. Journal of Applied Ecology 46:511-515 (pdf).


Babak, P. and He, F. 2009. A neutral model of edge effects. Theoretical Population Biology 75:76-83 (pdf).


Legendre, P., Mi, X.C., Ren, H.B., Ma, K.P., Sun, I.F., Yu, M.J. and He, F. 2009. Partitioning beta diversity in a subtropical broad-leaved forest of China. Ecology 90:663-674 (pdf).


Lin, K., Zhang, D.-Y. and He, F. 2009. Demographic trade-offs in a neutral model explain death rate-abundance rank relationship. Ecology 90:31-38 (pdf).


Chen, X.-Y. and He, F. 2009. Speciation and endemism under the model of island biogeography. Ecology 90:39-45 (pdf).


Li, L., Huang, Z.-L., Ye, W.-H., Cao, H.-L., Wei, S.-G., Wang, Z.-G., Lian, J.-Y., Sun, I.-F., Ma, K.-P. and He, F. 2009. Spatial patterns of tree species in a subtropical forest of China. Oikos 118:495-502 (pdf).


Hubbell, S.P., He, F., Condit, R., Borda-de-Água, L., Kellner, J. and ter Steege, H. 2008. How many tree species are there in the Amazon and how many of them will go extint? PNAS 105:11498-11504 (pdf).


Babak, P. and He, F. 2008. Species abundance distribution and dynamics in two locally coupled communities. Journal of Theoretical Biology 253:739-748 (pdf).

Gamarra, J.G.P. and He, F. 2008. Spatial scaling of mountain pine beetle infestations. Journal of Animal Ecology 77:796-801 (pdf) (Aber News;BBC news)
Zhu, H.T., He, F. and Zhou, J. 2008. Auto-multicategorical regression model for the distribution of vegetation. Statistics and Its Interface (pdf).
Getzin, S., Wiegand, T., Wiegand, K. and He, F. 2008. Heterogeneity influences spatial patterns and demographics in forest stands. Journal of Ecology 96:807-820 (pdf)
Shen, T.-J. and He, F. 2008. An incidence-based richness estimator for quadrats sampled without replacement. Ecology 89:2052-2060 (pdf).
Witte, J.-P. M., He, F. and Groen, C. 2008. Grid origin affects scaling of species across spatial scales. Global Ecology and Biogeography 17:448-456 (pdf).
He, F. and Tang, D.L. 2008. Estimating the niche preemption parameter of the geometric series. Acta Oecologica 33:105-107 (pdf).
Nelson, W., Potapov, A., Lewis, M., Hundsdorfer, A. and He, F. 2008. The balance of complexity in mechanistic modeling: Risk analysis in the mountain pine beetle. Journal of Applied Ecology 45:248-257 (pdf).
Pueyo, S., He, F. and Zillio, T. 2007. The maximum entropy formalism and the idiosyncratic theory of biodiversity. Ecology Letters 10:1017-1028 (pdf).
Hu, X.-S., Yeh, F.C. and He, F. 2007. Sex-ratio distortion driven by migration loads. Theoretical Population Biology 72:547-559 (pdf).
Hu, X.-S., He, F. and Hubbell, S.P. 2007. Species diversity in local neutral communities. American Naturalist 170:844-853 (pdf).
He, F. and Gaston, K. 2007. Estimating abundance from occurrence: an underdetermined problem. American Naturalist 170:655-659 (pdf).
McGill, B.J., Etienne, R.S., Gray, J.S., Alonso, D., Anderson , M.J., Benecha, H.K., Dornelas, M., Enquist, B.J., Green, J., He, F., Hurlbert, A., Magurran, A.E., Marquet, P.A., Maurer, B.A., Ostling, A., Soykan, C.U., Ugland, K., White, E. 2007. Species abundance distributions: moving beyond single prediction theories to integration within an ecological framework. Ecology Letters 10:995-1015 (pdf).
Getzin, S., Dean, C., He, F., Trofymow, T., Wiegand, K. and Wiegand, T. 2006. Spatial patterns and competition of tree species in a Douglas-fir chronosequence on Vancouver Island. Ecography 29:671-682 (pdf).
Hu, X.-S., He, F. and Hubbell, S.P. 2006. Neutral theory in population genetics and macroecology. Oikos 113:548-556 (pdf).
Magnussen, S., Pélissier, R., He, F. and Ramesh, B.R. 2006. An assessment of sample-based estimators of tree species richness in two wet tropical forest compartments in Panama and India. International Forestry Review 8:417-431 (pdf).
Gaston, K.J., Borges, P.A.V., He, F. and Gaspar, C. 2006. Abundance, spatial variance, and occupancy: species distribution in the Azores. Journal of Animal Ecology 75:646-656 (pdf) .
Wills, C., Harms, K.E., Condit, R., King, D., Thompson, J., He, F., Muller-Landau, H.C., Ashton, P., Losos, L., Comita, L., Hubbell, S.P., LaFrankie, J., Bunyavejchewin, P., Dattaraja, H.S., Davies, S., Esufali, S., Foster, R., Gunatilleke, N., Gunatilleke, S., Hall, P., Itoh, A., John, R., Kiratiprayoon, S., Loo de Lao, S., Massa, M., Nath, C., Noor, S., Kassim, A.R., Sukumar, R., Suresh, H.C., Sun, I.-F., Tan, S., Yamakura, T. and Zimmerman, J. 2006. Non-random processes contribute to the maintenance of diversity in tropical forests. Science 311:527-531 (pdf).
Hu, X.-S. and He, F. 2006. Seed and pollen flow in expanding a species' range. Journal of Theoretical Biology 240:662-672 (pdf).
Volkov, I., Banavar, J.R., He, F., Hubbell, S.P. and Maritan, A. 2006. Comparing models of species abundance: a reply. Nature 441:E1-E2 (pdf).
Muller-Landau, H.C., Condit, R.S., Harms, K.E., Marks, C.O., Thomas , S.C. , Bunyavejchewin, S., Chuyong, G., Co, L., Davies. S., Foster, R., Gunatilleke, S., Gunatilleke, N., Hart, T., Hubbell, S.P., Itoh, A., Kassim, A.R., Kenfack, D., LaFrankie, J.V., Lagunzad, D., Lee, H.S., Losos, E., Makana, J.-R., Ohkubo, T., Samper, C., Sukumar, R., Sun, I.-F., Nur Supardi M.N., Tan, S., Thomas, D., Thompson, J., Valencia, R., Vallejo, M.I., Muñoz, G.V., Yamakura, T. Zimmerman, J.K., Dattaraja, H.S., Esufali, S., Hall, P., He, F., Hernandez, C., Kiratiprayoon, S., Suresh, H.S., Wills, C. & Ashton, P. 2006. Comparing tropical forest tree size distributions with the predictions of metabolic ecology and equilibrium models. Ecology Letters 9:589-602.
He, F. and Hubbell, S.P. 2005. Percolation theory for the distribution and abundance of species: a reply. Physical Review Letters 95, article 189804 (pdf).
Volkov, I., Banavar, J.R., He, F., Hubbell, S.P. and Maritan, A. 2005. Density dependence explains tree species abundance and diversity in tropical forests. Nature 438:658-661 (pdf).
He, F. and Hu, X.-S. 2005. Hubbell's fundamental biodiversity parameter and the Simpson diversity index. Ecology Letters 8:386-390 (pdf).
Hu, X.-S. and He, F. 2005. Background selection and population differentiation. Journal of Theoretical Biology 235:207-219 (pdf).
He, F. 2005. Deriving a neutral model of species abundance from fundamental mechanisms of population dynamics. Functional Ecology 19:187-193 (pdf)
He, F., Gaston, K.J., Connor, E.F. and Srivastava, D. 2005. The local-regional relationship: immigration, extinction, and scale. Ecology 86:360-365 (pdf).
Wu. J., Huang, J.-H., Han, X.-G., Gao, X.-M., He, F., Jiang, M.-X., Jiang, Z.-G., Primack, R. B. and Shen, Z.-H. 2004. Three Georges Dam: An ecological perspective. Frontiers in Ecology and the Environment 2:241-248 (pdf).
He, F. and Hubbell, S.P. 2003. Percolation theory for the distribution and abundance of species. Physical Review Letters 91, article 198103 (pdf).
He, F. and Gaston, K.J. 2003. Occupancy, spatial variance, and the abundance of species. American Naturalist 162:366-375 (pdf).
Trofymow, J.A., Addison, J., Blackwell, B.A., He, F., Preston, C.A. and Marshall, V.G. 2003. Attributes and indicators of old-growth and successional Douglas-fir forests on Vancouver Island. Environmental Reviews 11:S187-S204.
He, F., Zhou, J. and Zhu, H.T. 2003. Autologistic regression model for the distribution of vegetation. Journal of Agricultural, Biological and Environmental Statistics 8:205-222 (pdf).
He, F., LaFrankie, J.V. and Song, B. 2002. Scale dependence of tree abundance and richness in a tropical rain forest, Malaysia. Landscape Ecology 17:559-568 (pdf).
Gaston, K.J. and He, F. 2002. The distribution of species range size: a stochastic process. Proceedings of the Royal Society of London, B 269:1079-1086 (pdf).
He, F. and Legendre, P. 2002. Species diversity patterns derived from species-area models. Ecology 83:1185-1198 (pdf).
He, F., Gaston, K.J. and Wu, J. 2002. On species occupancy-abundance models. ÉcoScience 9:119-126 (pdf).
Holt, A.R., Gaston, K.J. and He, F. 2002. Occupancy-abundance relationships and spatial distribution: a review. Basic & Applied Ecology 3:1-13.
He, F. and Gaston, K.J. 2000. Estimating species abundance from occurrence. American Naturalist 156:553-559 (pdf).
He, F. and Duncan, R.P. 2000. Density-dependent effects on tree survival in an old-growth Douglas-fir forest. Journal of Ecology 88:676-688 (pdf).
He, F. and Alfaro , R.I. 2000. White pine weevil attack on white spruce: a survival time analysis. Ecological Applications 10:35-42 (pdf).
He, F. and Gaston, K.J. 2000. Occupancy-abundance relationships and sampling scale. Ecography 23:503-511 (pdf).
He, F. and Barclay, H. 2000. Long-term response of understory plant species to thinning and fertilization in a Douglas-fir plantation on southern Vancouver Island , British Columbia. Canadian Journal of Forest Research 30:566-572 (pdf).
Tomlin, E.S., Alfaro , R.I., Borden, J.H. and He, F. 1998. Histological response of resistant and susceptible white spruce to simulated white pine weevil damage. Tree Physiology 18:21-28.
He, F., Legendre, P. and LaFrankie, J.V. 1997. Distribution patterns of tree species in a Malaysian tropical rain forest. Journal of Vegetation Science 8:105-114 (pdf).
Alfaro, R.I., He, F., Tomlin, E. and Kiss, G. 1997. White spruce resistance to white pine weevil related to bark resin canal density. Canadian Journal of Botany 75:568-573.
He, F. and Alfaro, R.I. 1997. White pine weevil (Coleoptera: Curculionidae) attack on white spruce: spatial and temporal patterns. Environmental Entomology 26:888-895 (pdf).
He, F. and Legendre, P. 1996. On species-area relations. American Naturalist 148:719-737 (pdf).
He, F., Legendre, P. and LaFrankie, J.V. 1996. Spatial patterning of diversity in a tropical rain forest in Malaysia. Journal of Biogeography 23:57-74 (pdf).
Alfaro, R.I., He, F., Kiss, G., King, J. and Yanchuk, A. 1996. Resistance of white spruce to white pine weevil: Development of a resistance index. Forest Ecology and Management 81:51-62.
Jeglum, J. and He, F. 1995. Pattern and vegetation-environment relationships in a boreal forested wetland in northeastern Ontario. Canadian Journal of Botany73:629-637.
He, F., Legendre, P., Belleheumer, C. and LaFrankie, J.V. 1994. Spatial scale and diversity patterns: a study of a tropical rain forest of Malaysia. Environmental and Ecological Statistics 1:265-286.
  Book Chapters
Gaston, K.J. and He. F. 2010. Species occurrence and occupancy. Pages 141-151 in Biological Diversity: Frontiers in Measurement and Assessment, eds. A. Maguran and B. McGill. Oxford University Press (pdf).
He, F. and Condit, R. 2007. The distribution of species: occupancy, scale and rarity. Pages 32-50 in Scaling Biodiversity, eds. D. Storch, P.L. Marquet and J.H. Brown. Cambridge University Press (pdf).
Borda de Agua, L., Hubbell, S.P. and He, F. 2007. Scaling biodiversity under neutrality. Pages 347-375 in Scaling Biodiversity, eds. D. Storch, P.L. Marquet and J.H. Brown. Cambridge University Press (pdf).
He, F. and Reed, W. 2006. Downscaling abundance from the distribution of species: occupancy theory and applications. Pages 89-108 in Scaling and Uncertainty Analysis in Ecology: Methods and Applications, eds. J. Wu, B. Jones, H. Li and O. Loucks. Springer, New York (pdf).
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