Latitudinal patterns of species range size of birds in China: A test of Rapoport’s rule

doi:10.17520/biods.2025274

Abstract

Abstract:

Aims: The variation of species distribution ranges along environmental gradients has long been an important topic in macroecology and biogeography. Rapoport’s rule was initially proposed as a hypothesis concerning the relationship between the latitudinal position of species distributions and their latitudinal range size. In 1996, Stevens provided a broad definition of Rapoport’s rule, stating that the geographic range size of species gradually increases with the rise of a biogeographic gradient (such as latitude, elevation, water depth, etc.). Currently, there is relatively limited research examining the applicability of Rapoport’s rule to latitudinal range breadth of Chinese birds, particularly in terms of comparisons across different taxonomic categories and traits.

Methods: We used distribution and trait data of Chinese birds, and applied four commonly used methods to test whether the latitudinal patterns of species range size of Chinese birds support Rapoport’s rule across three dimensions: taxonomic categories, feeding habits and body mass.

Results: The results showed that: (1) At different taxonomic levels (family, genus and species), higher taxonomic levels exhibited stronger support for Rapoport’s rule (the R² values for Stevens’s method, Pagel’s method and Cross-species method were all greater than 0.5). (2) Among different feeding habits, species range size variation of herbivorous birds exhibited the least support for Rapoport’s rule (Cross-species method: R²= 0.24). (3) Under the criteria of different body mass, small-bodied birds showed the strongest support for Rapoport’s rule (Cross-species method: R²= 0.42).

Conclusion: This study found that birds with different dimensions show varying applicability to Rapoport’s rule. The stronger applicability of small-bodied birds to Rapoport’s rule may be due to their weaker competitive abilities, rapid growth rates and better adaptability to changing environment; the lower applicability of herbivorous birds may be related to their correlation with the distribution of plant species. This study discussed the potential reasons for these differences, which helps to understand the formation and maintenance of biodiversity distribution patterns at a macroscale.

Key words: birds, body mass, feeding habit, Rapoport’s rule, taxonomic category

Shixian Li, Gang Feng. Latitudinal patterns of species range size of birds in China: A test of Rapoport’s rule[J]. Biodiv Sci, 2026, 34(3): 25274.

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URL: https://www.biodiversity-science.net/EN/10.17520/biods.2025274

https://www.biodiversity-science.net/EN/Y2026/V34/I3/25274

Figures/Tables 3

Fig. 1 Test of Rapoport’s rule for the different taxonomic categories of birds along the latitudinal gradients. Figure 1a, b, c, d showed the validation using Stevens’s method, Pagel’s method, Cross-species method and Mid-point method at the species level respectively; Figure 1e, f, g, h presented the validation using four methods at the genus level respectively; Figure 1i, j, k, l demonstrated the validation using four methods at the family level respectively. ** P < 0.01.

Fig. 2 Test of Rapoport’s rule for the different feeding habits of birds along the latitudinal gradients. Figure 2a, b, c, d showed the validation using Stevens’s method, Pagel’s method, Cross-species method and Mid-point method in carnivorous birds respectively; Figure 2e, f, g, h presented the validation using four methods in herbivorous birds respectively; Figure 2i, j, k, l demonstrated the validation using four methods in omnivorous birds respectively. ** P < 0.01.

Fig. 3 Test of Rapoport’s rule for the different body mass of birds along the latitudinal gradients. Figure 3 a, b, c, d showed the validation using Stevens’s method, Pagel’s method, Cross-species method and Mid-point method in small-bodied birds respectively; Figure 3 e, f, g, h presented the validation using four methods in large-bodied birds respectively. ** P < 0.01.

References 73

[1]	Alzate A, Rozzi R, Velasco JA, Robertson DR, Zizka A, Tobias JA, Hill A, Bacon CD, Janzen T, Pellissier L, van der Plas F, Rosindell J, Onstein RE (2025) Evolutionary age correlates with range size across plants and animals. Nature Communications, 16, 7894. DOI
[2]	Barnagaud JY, Kissling WD, Tsirogiannis C, Fisikopoulos V, Villéger S, Sekercioglu CH, Svenning JC (2017) Biogeographical, environmental and anthropogenic determinants of global patterns in bird taxonomic and trait turnover. Global Ecology and Biogeography, 26, 1190-1200. DOI URL
[3]	Barnagaud JY, Mazet N, Munoz F, Grenié M, Denelle P, Sobral M, Kissling WD, Şekercioğlu ÇH, Violle C (2019) Functional biogeography of dietary strategies in birds. Global Ecology and Biogeography, 28, 1004-1017. DOI URL
[4]	Barraclough TG, Humphreys AM (2015) The evolutionary reality of species and higher taxa in plants: A survey of post-modern opinion and evidence. New Phytologist, 207, 291-296. DOI PMID
[5]	Bateson W (1923) Age and area: A study in geographical distribution and origin of species. Nature, 111, 39-43. DOI
[6]	Blackburn TM, Gaston KJ (1994) The distribution of body sizes of the world’s bird species. Oikos, 70, 127-130. DOI URL
[7]	Boakes EH, McGowan PJK, Fuller RA, Ding CQ, Clark NE, O’Connor K, Mace GM (2010) Distorted views of biodiversity: Spatial and temporal bias in species occurrence data. PLoS Biology, 8, e1000385.
[8]	Bonino MF, Cruz FB (2025) Biological and environmental drivers of Liolaemus species distribution: A case story East of the Andes. Zoology, 173, 126303. DOI URL
[9]	Brown JH, Stevens GC, Kaufman DM (1996) The geographic range: Size, shape, boundaries, and internal structure. Annual Review of Ecology, Evolution, and Systematics, 27, 597-623.
[10]	Cardador L, De Cáceres M, Bota G, Giralt D, Casas F, Arroyo B, Mougeot F, Cantero-Martínez C, Moncunill J, Butler SJ, Brotons L (2014) A resource-based modelling framework to assess habitat suitability for steppe birds in semiarid Mediterranean agricultural systems. PLoS ONE, 9, e92790. DOI URL
[11]	Chen KY, Wang B, Chen C, Zhou GY (2024) The relationship between niche breadth and phylogenetic characteristics of eight species of rhubarb on the Qinghai-Tibet Plateau, Asia. Ecology and Evolution, 14, e11040. DOI URL
[12]	Chen W, Wang X, Cai YY, Huang XL, Li P, Liu W, Chang Q, Hu CC (2024) Potential distribution patterns and species richness of avifauna in rapidly urbanizing East China. Ecology and Evolution, 14, e11515. DOI URL
[13]	Chen Y, Wang JF, Zhang H, Ding HL, Tang SX (2012) Observation on feeding habit of dominant birds and forage sites in an evergreen broadleaved forest, Tiantong Forest Park, Zhejiang. Chinese Journal of Zoology, 47(5), 46-53. (in Chinese with English abstract)
	[陈宇, 王军馥, 张航, 丁虎林, 唐思贤 (2012) 天童常绿阔叶林样地中优势鸟类食性与采食地点植物的关系. 动物学杂志, 47(5), 46-53.]
[14]	De Bie T, De Meester L, Brendonck L, Martens K, Goddeeris B, Ercken D, Hampel H, Denys L, Vanhecke L, Van der Gucht K, Van Wichelen J, Vyverman W, Declerck SAJ (2012) Body size and dispersal mode as key traits determining metacommunity structure of aquatic organisms. Ecology Letters, 15, 740-747. DOI PMID
[15]	Ding JJ, Liu DZ, Li CW, Jiang ZG (2012) Spatial variation in species richness of birds and mammals in China’s mainland. Acta Ecologica Sinica, 32, 343-350. (in Chinese with English abstract) DOI URL
	[丁晶晶, 刘定震, 李春旺, 蒋志刚 (2012) 中国大陆鸟类和兽类物种多样性的空间变异. 生态学报, 32, 343-350.]
[16]	Gao W (2006) Studies on Birds and Their Ecology in Northeast China. Science Press, Beijing. (in Chinese)
	[高玮 (2006) 中国东北地区鸟类及其生态学研究. 科学出版社, 北京.]
[17]	Gaston KJ, Blackburn TM, Spicer JI (1998) Rapoport’s rule: Time for an epitaph? Trends in Ecology & Evolution, 13, 70-74. DOI URL
[18]	Guo QF, Qian H, Liu PC, Zhang J (2024) Macroecological correlates of richness, body size, and species range size in terrestrial vertebrates across the world. Frontiers of Biogeography, 16, e61729.
[19]	Hu JH, Hu HJ, Jiang ZG (2007) Distribution regularities of species diversity at large spatial scale. Chinese Journal of Applied & Environmental Biology, 13, 731-735. (in Chinese with English abstract)
	[胡军华, 胡慧建, 蒋志刚 (2007) 大空间尺度上物种多样性的分布规律. 应用与环境生物学报, 13, 731-735.]
[20]	Huang JH (1994) The spatial pattern of species diversity and its forming mechanism. Chinese Biodiversity, 2, 103-107. (in Chinese with English abstract)
	[黄建辉 (1994) 物种多样性的空间格局及其形成机制初探. 生物多样性, 2, 103-107.]
[21]	Inostroza-Michael O, Hernández CE, Rodríguez-Serrano E, Avaria-Llautureo J, Rivadeneira MM (2018) Interspecific geographic range size-body size relationship and the diversification dynamics of Neotropical furnariid birds. Evolution, 72, 1124-1133. DOI PMID
[22]	Jenkins CN, Pimm SL, Joppa LN (2013) Global patterns of terrestrial vertebrate diversity and conservation. Proceedings of the National Academy of Sciences, USA, 110, E2602-E2610.
[23]	Jin DM, Yang L, Xu ZP, Xiao C, Luo MF, Ma KP (2023) Effectiveness analysis of the National Specimen Information Infrastructure (NSII) in supporting scientific research on biodiversity. Guihaia, 43, 1501-1515. (in Chinese with English abstract)
	[金冬梅, 杨灵, 许哲平, 肖翠, 罗茂芳, 马克平 (2023) 国家标本资源共享平台(NSII)支撑生物多样性科学研究的成效分析. 广西植物, 43, 1501-1515.]
[24]	Laube I, Korntheuer H, Schwager M, Trautmann S, Rahbek C, Böhning-Gaese K (2013) Towards a more mechanistic understanding of traits and range sizes. Global Ecology and Biogeography, 22, 233-241. DOI URL
[25]	Lawrence WS (1987) Dispersal: An alternative mating tactic conditional on sex ratio and body size. Behavioral Ecology and Sociobiology, 21, 367-373. DOI URL
[26]	Lei FM, Song G, Cai TL, Qu YH, Jia CX, Zhao YF, Zhang DZ (2021) Research progress and prospect on biogeography of birds in China. Chinese Journal of Zoology, 56, 265-289. (in Chinese with English abstract)
	[雷富民, 宋刚, 蔡天龙, 屈延华, 贾陈喜, 赵义方, 张德志 (2021) 中国鸟类生物地理学研究回顾与展望. 动物学杂志, 56, 265-289.]
[27]	Letcher AJ, Harvey PH (1994) Variation in geographical range size among mammals of the Palearctic. The American Naturalist, 144, 30-42. DOI URL
[28]	Li DL, Zhang J, Liu Y, Lloyd H, Pagani-Núñez E, Zhang ZW (2020) Differences in dietary specialization, habitat use and susceptibility to human disturbance influence feeding rates and resource partitioning between two migratory Numenius curlew species. Estuarine, Coastal and Shelf Science, 245, 106990. DOI URL
[29]	Li LP, He SY, Jiang YM, Wang T, Zhao HH, Cui WH, Zheng YM, Hai Y, Wan HW (2019) Species range size patterns and their significance on biodiversity conservation. Scientia Sinica (Vitae), 49, 929-937. (in Chinese with English abstract)
	[李利平, 何思源, 蒋样明, 王拓, 赵辉辉, 崔伟宏, 郑姚闽, 海鹰, 万华伟 (2019) 物种分布区特征及其对生物多样性保育的意义. 中国科学: 生命科学, 49, 929-937.]
[30]	Li MS, Wu XQ, Zhou YJ, Long SY, Zhang MH, Gu HW, Si XF, Yan ER, Zhang J (2024) Elevational pattern of species range size of vascular plants in Changbai Mountain, China: A test of Rapoport’s rule. Chinese Journal of Applied Ecology, 35, 3329-3338. (in Chinese with English abstract)
	[李漫淑, 吴晓晴, 周银佳, 龙诗怡, 张美惠, 顾汉威, 斯幸峰, 阎恩荣, 张健 (2024) 长白山维管植物海拔梯度上种域格局及Rapoport法则的检验. 应用生态学报, 35, 3329-3338.] DOI
[31]	Li YQ, Wang ZH (2023) Functional biogeography of plants: Research progresses and challenges. Chinese Journal of Plant Ecology, 47, 145-169. (in Chinese with English abstract) DOI URL
	[李耀琪, 王志恒 (2023) 植物功能生物地理学的研究进展与展望. 植物生态学报, 47, 145-169.] DOI
[32]	Liang HZ, Fu TG, Gao H, Li M, Liu JT (2023) Climatic and non-climatic drivers of plant diversity along an altitudinal gradient in the Taihang Mountains of northern China. Diversity, 15, 66. DOI URL
[33]	Liu C, Zheng CY, Zhang T, Zeng FX, Wang YR (2014) Geographic patterns of avian species richness in China and their environmental factors. Acta Scientiarum Naturalium Universitatis Pekinensis, 50, 429-438. (in Chinese with English abstract)
	[刘澈, 郑成洋, 张腾, 曾发旭, 王逸然 (2014) 中国鸟类物种丰富度的地理格局及其与环境因子的关系. 北京大学学报(自然科学版), 50, 429-438.]
[34]	Liu XY, Li JS, Zhao CY, Quan ZJ, Zhao XJ, Gong L (2016) Prediction of potential suitable area of Ambrosia artemisiifolia L. in China based on MAXENT and ArcGIS. Journal of Plant Protection, 43, 1041-1048. (in Chinese with English abstract)
	[柳晓燕, 李俊生, 赵彩云, 全占军, 赵相健, 宫璐 (2016) 基于MAXENT模型和ArcGIS预测豚草在中国的潜在适生区. 植物保护学报, 43, 1041-1048.]
[35]	Luan L, Jiang YJ, Cheng MH, Dini-Andreote F, Sui YY, Xu QS, Geisen S, Sun B (2020) Organism body size structures the soil microbial and nematode community assembly at a continental and global scale. Nature Communications, 11, 6406. DOI PMID
[36]	Lustenhouwer N, Riddell EA (2025) Range shifts as drivers of niche breadth and dispersal ability in wild populations. Journal of Animal Ecology, 94, 175-178. DOI PMID
[37]	Maurer BA (1998) The evolution of body size in birds. II. The role of reproductive power. Evolutionary Ecology, 12, 935-944. DOI URL
[38]	McCain CM, Knight BK (2013) Elevational Rapoport’s rule is not pervasive on mountains. Global Ecology and Biogeography, 22, 750-759. DOI URL
[39]	Morales-Castilla I, Rodríguez MÁ, Kaur R, Hawkins BA (2013) Range size patterns of New World oscine passerines (Aves): Insights from differences among migratory and sedentary clades. Journal of Biogeography, 40, 2261-2273. DOI URL
[40]	Nilsson JA (1989) Causes and consequences of natal dispersal in the marsh tit, Parus palustris. Journal of Animal Ecology, 58, 619-636. DOI URL
[41]	Orkney A, Hedrick BP (2024) Small body size is associated with increased evolutionary lability of wing skeleton proportions in birds. Nature Communications, 15, 4208. DOI PMID
[42]	Pagel MD, May RM, Collie AR (1991) Ecological aspects of the geographical distribution and diversity of mammalian species. The American Naturalist, 137, 791-815. DOI URL
[43]	Qian H, Ricklefs RE (2004) Taxon richness and climate in angiosperms: Is there a globally consistent relationship that precludes region effects? The American Naturalist, 163, 773-779. PMID
[44]	R Core Team (2022) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria.
[45]	Rapoport EH (1975) Areografía: Estrategias geográficas de las especies. Fondo de Cultura Económica, Mexico City.
[46]	Read QD, Baiser B, Grady JM, Zarnetske PL, Record S, Belmaker J (2018) Tropical bird species have less variable body sizes. Biology Letters, 14, 20170453. DOI URL
[47]	Reif J, Hořák D, Krištín A, Kopsová L, Devictor V (2016) Linking habitat specialization with species’ traits in European birds. Oikos, 125, 405-413. DOI URL
[48]	Rohde K (1992) Latitudinal gradients in species diversity: The search for the primary cause. Oikos, 65, 514-527. DOI URL
[49]	Saino N, Romano M, Scandolara C, Rubolini D, Ambrosini R, Caprioli M, Costanzo A, Romano A (2014) Brownish, small and lousy barn swallows have greater natal dispersal propensity. Animal Behaviour, 87, 137-146. DOI URL
[50]	Saupe EE, Myers CE, Peterson AT, Soberón J, Singarayer J, Valdes P, Qiao HJ (2019) Non-random latitudinal gradients in range size and niche breadth predicted by spatial patterns of climate. Global Ecology and Biogeography, 28, 928-942. DOI
[51]	Sax DF (2001) Latitudinal gradients and geographic ranges of exotic species: Implications for biogeography. Journal of Biogeography, 28, 139-150. DOI URL
[52]	Seifert CL, Strutzenberger P, Hausmann A, Fiedler K (2022) Dietary specialization mirrors Rapoport’s rule in European geometrid moths. Global Ecology and Biogeography, 31, 1161-1171. DOI URL
[53]	Sheard C, Neate-Clegg MHC, Alioravainen N, Jones SEI, Vincent C, MacGregor HEA, Bregman TP, Claramunt S, Tobias JA (2020) Ecological drivers of global gradients in avian dispersal inferred from wing morphology. Nature Communications, 11, 2463. DOI PMID
[54]	Shen ZH, Lu QY (2009) The Rapoport’s rule for the geographic patterns of species range size. Biodiversity Science, 17, 560-567. (in Chinese with English abstract) DOI URL
	[沈泽昊, 卢绮妍 (2009) 物种分布区范围地理格局的Rapoport法则. 生物多样性, 17, 560-567.] DOI
[55]	Si XF, Ding P (2011) History, status of monitoring land birds in Europe and America and countermeasures of China. Biodiversity Science, 19, 303-310. (in Chinese with English abstract) DOI URL
	[斯幸峰, 丁平 (2011) 欧美陆地鸟类监测的历史、现状与我国的对策. 生物多样性, 19, 303-310.] DOI
[56]	Slatyer RA, Hirst M, Sexton JP (2013) Niche breadth predicts geographical range size: A general ecological pattern. Ecology Letters, 16, 1104-1114. DOI PMID
[57]	Stevens GC (1989) The latitudinal gradient in geographical range: How so many species coexist in the tropics. The American Naturalist, 133, 240-256. DOI URL
[58]	Stevens GC (1996) Extending Rapoport’s rule to Pacific marine fishes. Journal of Biogeography, 23, 149-154. DOI URL
[59]	Sun W, Pan XY, Liang JC, Ding ZF, Zhou J, Hu HJ (2021) Species range pattern of breeding birds in the middle part of the Himalayas with a test of Rapoport’s rule—Based on data from an altitude gradient of 3600 m. Chinese Journal of Zoology, 56, 358-366. (in Chinese with English abstract)
	[孙雯, 潘新园, 梁健超, 丁志锋, 周江, 胡慧建 (2021) 喜马拉雅中段繁殖鸟类种域格局及Rapoport法则检验——基于3600 m海拔梯度范围的数据. 动物学杂志, 56, 358-366.]
[60]	Wang J, Gao ZZ, Jiang YT, Wan DM (2021) Research advances in the intestinal microbes in wild birds with different feeding habits. Acta Ecologica Sinica, 41, 7939-7945. (in Chinese with English abstract)
	[王娟, 高泽中, 蒋一婷, 万冬梅 (2021) 不同食性野生鸟类肠道微生物研究进展. 生态学报, 41, 7939-7945.]
[61]	Wang YP, Song YF, Zhong YX, Chen CW, Zhao YH, Zeng D, Wu YR, Ding P (2021) A dataset on the life-history and ecological traits of Chinese birds. Biodiversity Science, 29, 1149-1153. (in Chinese with English abstract) DOI URL
	[王彦平, 宋云枫, 钟雨茜, 陈传武, 赵郁豪, 曾頔, 吴亦如, 丁平 (2021) 中国鸟类的生活史和生态学特征数据集. 生物多样性, 29, 1149-1153.] DOI
[62]	Weiss-Lehman C, Shaw AK (2022) Understanding the drivers of dispersal evolution in range expansions and their ecological consequences. Evolutionary Ecology, 36, 181-197. DOI
[63]	Willig MR, Kaufman DM, Stevens RD (2003) Latitudinal gradients of biodiversity: Pattern, process, scale, and synthesis. Annual Review of Ecology, Evolution, and Systematics, 34, 273-309. DOI URL
[64]	Ye YX, Santoro S, Song ZT, Hu CS, Zhang Z, Qing BP, Wang C, Ding CQ (2023) Dispersal patterns of the endangered Crested Ibis suggest high breeding densities drive natal dispersal. Ornithological Applications, 125, duac042. DOI URL
[65]	Zhang WJ, Lu QY, Liang J, Shen ZH (2010) Altitudinal gradients of species richness and range size of vascular plants in Taiwan: A test of Rapoport’s rule. Biodiversity Science, 18, 312-322. (in Chinese with English abstract) DOI URL
	[张婉君, 卢绮妍, 梁军, 沈泽昊 (2010) 台湾维管束植物物种丰富度和种域宽度的海拔格局及对Rapoport法则的检验. 生物多样性, 18, 312-322.] DOI
[66]	Zheng GM (2017) A Checklist on the Classification and Distribution of the Birds of China. Science Press, Beijing. (in Chinese)
	[郑光美 (2017) 中国鸟类分类与分布名录. 科学出版社, 北京.]
[67]	Zheng GM (2023) A Checklist on the Classification and Distribution of the Birds of China, 4th edn. Science Press, Beijing. (in Chinese)
	[郑光美 (2023) 中国鸟类分类与分布名录(第四版). 科学出版社, 北京.]
[68]	Zheng Z, Gong DJ, Sun CX (2014a) Elevational pattern of species richness and species range size of herpetofauna in Baishuijiang Nature Reserve: A test of Rapoport’s rule. Chinese Journal of Ecology, 33, 537-546. (in Chinese with English abstract)
	[郑智, 龚大洁, 孙呈祥 (2014a) 白水江自然保护区两栖爬行动物物种丰富度和种域海拔梯度格局及对Rapoport法则的验证. 生态学杂志, 33, 537-546.]
[69]	Zheng Z, Gong DJ, Sun CX, Li XJ, Li WJ (2014b) Altitudinal patterns of species richness and species range size of vascular plants in Xiaolongshan Reserve of Qinling Mountain: A test of Rapoport’s rule. Chinese Journal of Applied Ecology, 25, 2477-2485. (in Chinese with English abstract)
	[郑智, 龚大洁, 孙呈祥, 李晓军, 李万江 (2014b) 秦岭小陇山保护区维管植物丰富度和种域海拔梯度格局及其对Rapoport法则验证. 应用生态学报, 25, 2477-2485.]
[70]	Zhi YJ, Shao MQ, Li QJ (2020) Wading bird habitat, water depth utilization and niche separation in Poyang Lake, China. Pakistan Journal of Zoology, 52, 2243-2250.
[71]	Zhu C, Dalsgaard B, Li WD, Kaiser-Bunbury CN, Simmons BI, Ren P, Zhao YH, Zeng D, Gonçalves F, Zhang X, Chang LX, Ding P, Si XF (2025) Interconnecting fragmented forests: Small and mobile birds are cornerstones in the plant-frugivore meta-network. Proceedings of the National Academy of Sciences, USA, 122, e2415846122.
[72]	Zimova M, Weeks BC, Willard DE, Giery ST, Jirinec V, Burner RC, Winger BM (2023) Body size predicts the rate of contemporary morphological change in birds. Proceedings of the National Academy of Sciences, USA, 120, e2206971120.
[73]	Zou FS, Ye GF (2016) Distribution List of Terrestrial Vertebrates in Guangdong. Guangdong Science and Technology Press, Guangzhou. (in Chinese)
	[邹发生, 叶冠锋 (2016) 广东陆生脊椎动物分布名录. 广东科技出版社, 广州.]