Species packing and the latitudinal gradient in beta-diversity

Ke Cao, Richard Condit, Xiangcheng Mi, Lei Chen, Haibao Ren, Wubing Xu, David F R P Burslem, Chunrong Cai, Min Cao, Li-Wan Chang, Chengjin Chu, Fuxin Cui, Hu Du, Sisira Ediriweera, C S V Gunatilleke, I U A N Gunatilleke, Zhanqing Hao, Guangze Jin, Jinbo Li, Buhang LiYide Li, Yankun Liu, Hongwei Ni, Michael J O'Brien, Xiujuan Qiao, Guochun Shen, Songyan Tian, Xihua Wang, Han Xu, Yaozhan Xu, Libing Yang, Sandra L Yap, Juyu Lian, Wanhui Ye, Mingjian Yu, Sheng-Hsin Su, Chia-Hao Chang-Yang, Yili Guo, Xiankun Li, Fuping Zeng, Daoguang Zhu, Li Zhu, I-Fang Sun, Keping Ma, Jens-Christian Svenning

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The decline in species richness at higher latitudes is among the most fundamental patterns in ecology. Whether changes in species composition across space (beta-diversity) contribute to this gradient of overall species richness (gamma-diversity) remains hotly debated. Previous studies that failed to resolve the issue suffered from a well-known tendency for small samples in areas with high gamma-diversity to have inflated measures of beta-diversity. Here, we provide a novel analytical test, using beta-diversity metrics that correct the gamma-diversity and sampling biases, to compare beta-diversity and species packing across a latitudinal gradient in tree species richness of 21 large forest plots along a large environmental gradient in East Asia. We demonstrate that after accounting for topography and correcting the gamma-diversity bias, tropical forests still have higher beta-diversity than temperate analogues. This suggests that beta-diversity contributes to the latitudinal species richness gradient as a component of gamma-diversity. Moreover, both niche specialization and niche marginality (a measure of niche spacing along an environmental gradient) also increase towards the equator, after controlling for the effect of topographical heterogeneity. This supports the joint importance of tighter species packing and larger niche space in tropical forests while also demonstrating the importance of local processes in controlling beta-diversity.

Original languageEnglish
Article number20203045
Number of pages8
JournalProceedings of the Royal Society of London. B, Biological Sciences
Issue number1948
Early online date14 Apr 2021
Publication statusPublished - 14 Apr 2021

Bibliographical note


We thank Dingliang Xing, Tak Fung, Fangliang He and Gabriel Arellano for comments on the earlier draft. We thank Alex Karolus for leading the census in the Danum Valley forest plot, and we are grateful to Mike Bernados and Bill McDonald for species identifications, to Fangliang He, Stuart Davies and Shameema Esufali for advice and training, to Qianjiangyuan National Park, the Center for Forest Science at Morton Arboretum, Fushan Research Center, Lienhuachih Research Center and Sri Lankan Forest Department for logistical support and the hundreds of fieldworkers and students who measured and mapped the trees analysed in this study.

Funding. This work was financially supported by the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB31000000) and National Natural Science Foundation of China (NSFC 31770478). Data collection was funded by many organizations,principally, NSFC 31470490, 31470487, 41475123, 31570426, 31570432, 31570486, 31622014, 31660130, 31670441, 31670628, 31700356, 31760141, 31870404 and 32061123003, the Southeast Asia Rain Forest Research Programme (SEARRP), National Key Basic Research Program of China (Grant No. 2014CB954100), SEARRP partners especially Yayasan Sabah, HSBC Malaysia, financial project of Heilongjiang Pro- vince (XKLY2018ZR01), National Key R&D Program of China (2016YFC1201102 and 2016YFC0502405), the Central Public-interest Scientific Institution Basal Research Fund (CAFYBB2017ZE001), CTFS Forest GEO for funding for Sinharaja forest plot, the Taiwan For- estry Bureau (92-00-2-06 and tfbm960226), the Taiwan Forestry Research Institute (93AS-2.4.2-FI-G1, 94AS-11.1.2-FI-G1, and 97AS- 7.1.1.F1-G1) and the Ministry of Science and Technology of Taiwan (NSC92-3114-B002-009) for funding the Fushan and Lienhuachih plots, Scientific Research Funds of Heilongjiang Provincial Research Institutes (CZKYF2021B006). J.C.S. considers this work a contribution to his VILLUM Investigator project ‘Biodiversity Dynamics in a Changing World’ funded by VILLUM FONDEN (grant no. 16549).


  • sampling bias
  • niche specialisation
  • latitude
  • beta-diversity
  • gamma-diversity
  • species packing


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