Genetic diversity and genetic structure of Zhongshan partridge duck populations

doi:10.17520/biods.2024592

Abstract

Abstract:

Aims: The Zhongshan partridge duck is the only local duck breed in Guangdong Province and represents the most representative rare genetic resource in Zhongshan City. However, current research lacks a comprehensive understanding of the genetic diversity and genetic structure of Zhongshan partridge duck populations, limiting protection and resource utilization. The primary objectives of this study are to systematically analyze the genetic diversity and population structure of the Zhongshan partridge duck populations using whole-genome resequencing technology, evaluate the conservation effectiveness of its captive breeding programs in maintaining genetic integrity, and propose scientifically informed strategies for the protection of this genetic resource.

Methods: We collected 91 samples from seven duck populations in Guangdong and Guangxi provinces and obtained resequencing data for 388 samples from 19 additional populations through the NCBI database, generating a combined dataset of 479 individuals representing 22 duck breeds. Population-level variant detection was conducted using the GenomicsDB method in GATK software. Genetic diversity parameters—including effective number of alleles (Ne), observed heterozygosity (Ho), expected heterozygosity (He), polymorphism information content (PIC), and nucleotide diversity (π)—were systematically calculated for all 22 populations using VCFtools. Population genetic structure was evaluated through neighbor-joining (NJ) tree construction, principal component analysis (PCA), and Bayesian clustering analysis. Gene flow dynamics between the Zhongshan partridge duck and other breeds were subsequently investigated using TreeMix analysis.

Results: Genetic diversity assessments revealed that the Zhongshan partridge duck population exhibited relatively elevated genetic diversity among the 22 analyzed breeds (Ne = 3.8656, Ho = 0.3382, He = 0.3313). Notably, its Ho ranked highest across all breeds, while its He occupied the seventh position, still reflecting a substantial level of genetic variation. Population genetic structure analyses, including phylogenetic tree reconstruction and principal component analysis (PCA), demonstrated close clustering of the Zhongshan partridge duck with the Beijing Duck, Wenshi partridge duck, and maple leaf duck. Admixture analysis identified the Zhongshan partridge duck as a distinct genetic cluster characterized by a single ancestral component at K = 10. TreeMix analysis with Zhongshan partridge duck designated as the outgroup failed to detect significant gene flow from other breeds into this population. Analyses utilizing the mallard (Anas platyrhynchos) as the outgroup revealed only one migration event associated with the Zhongshan partridge duck, originating from the ancestral node of the Yulin partridge duck prior to its divergence.

Conclusion: Population fluctuations constitute a critical determinant of genetic diversity in avian conservation genetics. Despite undergoing historical demographic bottlenecks characterized by severe population contractions, the Zhongshan partridge duck retains comparatively elevated genomic diversity, a conservation outcome attributable to coordinated ex situ management protocols implemented through the collaborative efforts of the Zhongshan Agricultural Technology Extension Center and South China Agricultural University. Population genomic structure analyses substantiate the conservation efficacy of the Wugui Mountain Conservation Farm, demonstrating successful maintenance of ancestral allelic variation. This investigation delineates the extant genetic architecture and population stratification patterns of the Zhongshan partridge duck, formulates empirically grounded strategies for conserving its evolutionary significant units (ESUs), and establishes a genomic-informed paradigm for safeguarding this agriculturally vital germplasm resource.

Key words: genetic diversity, population genetic structure, Zhongshan partridge duck, gene flow analysis, whole-genome resequencing

Ruixiang Xue, Xuerong Ma, Jiongwen Wu, Aijun Liu, Xiquan Zhang, Congliang Ji, Yingshan Yin, Weijian Zhu, Qingbin Luo. Genetic diversity and genetic structure of Zhongshan partridge duck populations[J]. Biodiv Sci, 2025, 33(8): 24592.

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URL: https://www.biodiversity-science.net/EN/10.17520/biods.2024592

https://www.biodiversity-science.net/EN/Y2025/V33/I8/24592

Figures/Tables 5

Table 1 Genetic parameter table of population for 22 duck species/varieties

种类 Variety	有效等位基因数 Ne	观测杂合度 Ho	期望杂合度 He	多态信息含量 PIC	核苷酸多样 π
北京鸭 BJ	3.9826 ± 2.3280	0.2633 ± 0.1229	0.3221 ± 0.1335	0.2645 ± 0.0918	0.0030 ± 0.0024
斑嘴鸭 BZ	5.9716 ± 3.3542	0.1658 ± 0.1304	0.2389 ± 0.1392	0.2129 ± 0.1072	0.0034 ± 0.0028
东栏鸭 DL	4.1970 ± 2.6863	0.3091 ± 0.1863	0.3201 ± 0.1394	0.2938 ± 0.1224	0.0028 ± 0.0024
奉化鸭 FH	5.3629 ± 3.2577	0.1780 ± 0.1393	0.2661 ± 0.1437	0.2333 ± 0.1109	0.0028 ± 0.0023
枫叶鸭 FY	4.1817 ± 2.4431	0.2789 ± 0.1824	0.3138 ± 0.1376	0.2865 ± 0.1201	0.0020 ± 0.0020
广西小麻鸭 GX	4.2470 ± 2.6607	0.2902 ± 0.1591	0.3135 ± 0.1379	0.2708 ± 0.1035	0.0033 ± 0.0028
高邮鸭 GY	3.6321 ± 2.1210	0.2597 ± 0.1479	0.3399 ± 0.1251	0.2908 ± 0.0896	0.0018 ± 0.0015
金定鸭 JD	4.4242 ± 2.3207	0.1713 ± 0.1040	0.2865 ± 0.1272	0.2400 ± 0.0925	0.0001 ± 0.0001
靖西大麻鸭 JX	3.6073 ± 1.8445	0.3134 ± 0.1512	0.3364 ± 0.1254	0.2881 ± 0.0915	0.0032 ± 0.0027
连城白鸭 LC	4.6378 ± 2.3813	0.1771 ± 0.1074	0.2745 ± 0.1266	0.2317 ± 0.0926	0.0001 ± 0.0001
龙胜翠鸭 LS	3.5011 ± 1.6571	0.3160 ± 0.1638	0.3407 ± 0.1234	0.2995 ± 0.0948	0.0033 ± 0.0026
莆田黑鸭 PT	5.1533 ± 2.5464	0.1403 ± 0.1042	0.2495 ± 0.1241	0.2137 ± 0.0925	0.0001 ± 0.0001
融水香鸭 RS	4.5350 ± 3.0489	0.2930 ± 0.1835	0.3084 ± 0.1440	0.2803 ± 0.1242	0.0029 ± 0.0026
山麻鸭 SM	4.3646 ± 2.2142	0.1747 ± 0.0953	0.2863 ± 0.1241	0.2393 ± 0.0888	0.0001 ± 0.0001
绍兴鸭 SX	3.7232 ± 2.1620	0.2186 ± 0.1196	0.3358 ± 0.1284	0.2750 ± 0.0873	0.0020 ± 0.0016
温氏麻鸭 WMA	4.7271 ± 3.1733	0.3004 ± 0.1869	0.3009 ± 0.1460	0.2777 ± 0.1306	0.0034 ± 0.0029
文桥鸭 WQ	4.4776 ± 3.0149	0.2976 ± 0.1829	0.3108 ± 0.1432	0.2826 ± 0.1231	0.0032 ± 0.0027
温氏白鸭 WSB	4.3264 ± 2.9341	0.3048 ± 0.1847	0.3180 ± 0.1416	0.2897 ± 0.1266	0.0029 ± 0.0027
玉林麻鸭 YL	4.6123 ± 3.1255	0.2916 ± 0.1832	0.3070 ± 0.1460	0.2781 ± 0.1248	0.0030 ± 0.0026
樱桃谷鸭 YTG	4.0367 ± 2.3784	0.2378 ± 0.1587	0.3221 ± 0.1361	0.2799 ± 0.1034	0.0014 ± 0.0012
绿头野鸭 YY	4.5105 ± 2.5598	0.2350 ± 0.1284	0.2922 ± 0.1364	0.2437 ± 0.0954	0.0037 ± 0.0027
中山麻鸭 ZS	3.8656 ± 2.3322	0.3382 ± 0.1609	0.3313 ± 0.1329	0.2849 ± 0.1035	0.0032 ± 0.0029

Table 1 Genetic parameter table of population for 22 duck species/varieties

种类 Variety	有效等位基因数 Ne	观测杂合度 Ho	期望杂合度 He	多态信息含量 PIC	核苷酸多样 π
北京鸭 BJ	3.9826 ± 2.3280	0.2633 ± 0.1229	0.3221 ± 0.1335	0.2645 ± 0.0918	0.0030 ± 0.0024
斑嘴鸭 BZ	5.9716 ± 3.3542	0.1658 ± 0.1304	0.2389 ± 0.1392	0.2129 ± 0.1072	0.0034 ± 0.0028
东栏鸭 DL	4.1970 ± 2.6863	0.3091 ± 0.1863	0.3201 ± 0.1394	0.2938 ± 0.1224	0.0028 ± 0.0024
奉化鸭 FH	5.3629 ± 3.2577	0.1780 ± 0.1393	0.2661 ± 0.1437	0.2333 ± 0.1109	0.0028 ± 0.0023
枫叶鸭 FY	4.1817 ± 2.4431	0.2789 ± 0.1824	0.3138 ± 0.1376	0.2865 ± 0.1201	0.0020 ± 0.0020
广西小麻鸭 GX	4.2470 ± 2.6607	0.2902 ± 0.1591	0.3135 ± 0.1379	0.2708 ± 0.1035	0.0033 ± 0.0028
高邮鸭 GY	3.6321 ± 2.1210	0.2597 ± 0.1479	0.3399 ± 0.1251	0.2908 ± 0.0896	0.0018 ± 0.0015
金定鸭 JD	4.4242 ± 2.3207	0.1713 ± 0.1040	0.2865 ± 0.1272	0.2400 ± 0.0925	0.0001 ± 0.0001
靖西大麻鸭 JX	3.6073 ± 1.8445	0.3134 ± 0.1512	0.3364 ± 0.1254	0.2881 ± 0.0915	0.0032 ± 0.0027
连城白鸭 LC	4.6378 ± 2.3813	0.1771 ± 0.1074	0.2745 ± 0.1266	0.2317 ± 0.0926	0.0001 ± 0.0001
龙胜翠鸭 LS	3.5011 ± 1.6571	0.3160 ± 0.1638	0.3407 ± 0.1234	0.2995 ± 0.0948	0.0033 ± 0.0026
莆田黑鸭 PT	5.1533 ± 2.5464	0.1403 ± 0.1042	0.2495 ± 0.1241	0.2137 ± 0.0925	0.0001 ± 0.0001
融水香鸭 RS	4.5350 ± 3.0489	0.2930 ± 0.1835	0.3084 ± 0.1440	0.2803 ± 0.1242	0.0029 ± 0.0026
山麻鸭 SM	4.3646 ± 2.2142	0.1747 ± 0.0953	0.2863 ± 0.1241	0.2393 ± 0.0888	0.0001 ± 0.0001
绍兴鸭 SX	3.7232 ± 2.1620	0.2186 ± 0.1196	0.3358 ± 0.1284	0.2750 ± 0.0873	0.0020 ± 0.0016
温氏麻鸭 WMA	4.7271 ± 3.1733	0.3004 ± 0.1869	0.3009 ± 0.1460	0.2777 ± 0.1306	0.0034 ± 0.0029
文桥鸭 WQ	4.4776 ± 3.0149	0.2976 ± 0.1829	0.3108 ± 0.1432	0.2826 ± 0.1231	0.0032 ± 0.0027
温氏白鸭 WSB	4.3264 ± 2.9341	0.3048 ± 0.1847	0.3180 ± 0.1416	0.2897 ± 0.1266	0.0029 ± 0.0027
玉林麻鸭 YL	4.6123 ± 3.1255	0.2916 ± 0.1832	0.3070 ± 0.1460	0.2781 ± 0.1248	0.0030 ± 0.0026
樱桃谷鸭 YTG	4.0367 ± 2.3784	0.2378 ± 0.1587	0.3221 ± 0.1361	0.2799 ± 0.1034	0.0014 ± 0.0012
绿头野鸭 YY	4.5105 ± 2.5598	0.2350 ± 0.1284	0.2922 ± 0.1364	0.2437 ± 0.0954	0.0037 ± 0.0027
中山麻鸭 ZS	3.8656 ± 2.3322	0.3382 ± 0.1609	0.3313 ± 0.1329	0.2849 ± 0.1035	0.0032 ± 0.0029

Fig. 1 Phylogenetic tree of 479 duck individuals. ZS, Zhongshan partridge duck; BJ, Beijing duck; LC, Liancheng white duck; WMA, Wenshi partridge duck; WSB, Wenshi white duck; GX, Guangxi small partridge duck; JX, Jingxi large partridge duck; YY, Mallard (Anas platyrhynchos); BZ, Spot-billed duck (Anas poecilorhyncha); FY, Maple leaf duck; YTG, Cherry valley duck; SX, Shaoxing duck; FH, Fenghua duck; GY, Gaoyou duck; SM, Mountain partridge duck; JD, Jinding duck; PT, Putian black duck; LS, Longsheng emerald duck; DL, Donglan duck; RS, Rongshui Xiang duck; WQ, Wenqiao duck; YL, Yulin partridge duck.

Fig. 2 Principal component analysis (PCA) plot of 479 duck individuals based on PC1 and PC2. Letters in the diagram represent abbreviations of duck breed names (ZS, Zhongshan partridge duck; BJ, Beijing duck; LC, Liancheng white duck; WMA, Wenshi partridge duck; WSB, Wenshi white duck; GX, Guangxi small partridge duck; JX, Jingxi large partridge duck; YY, Mallard (Anas platyrhynchos); BZ, Spot-billed duck (Anas poecilorhyncha); FY, Maple leaf duck; YTG, Cherry valley duck; SX, Shaoxing duck; FH, Fenghua duck; GY, Gaoyou duck; SM, Mountain partridge duck; JD, Jinding duck; PT, Putian black duck; LS, Longsheng emerald duck; DL, Donglan duck; RS, Rongshui Xiang duck; WQ, Wenqiao duck; YL, Yulin partridge duck).

Fig. 3 Admixture analysis plot of 21 duck breeds from K = 2 to K = 10. ZS, Zhongshan partridge duck; BJ, Beijing duck; LC, Liancheng white duck; WMA, Wenshi partridge duck; WSB, Wenshi white duck; GX, Guangxi small partridge duck; JX, Jingxi large partridge duck; YY, Mallard (Anas platyrhynchos); BZ, Spot-billed duck (Anas poecilorhyncha); FY, Maple leaf duck; YTG, Cherry valley duck; SX, Shaoxing duck; FH, Fenghua duck; GY, Gaoyou duck; SM, Mountain partridge duck; JD, Jinding duck; PT, Putian black duck; LS, Longsheng emerald duck; DL, Donglan duck; RS, Rongshui Xiang duck; WQ, Wenqiao duck; YL, Yulin partridge duck.

Fig. 4 TreeMix analysis plots of 22 duck breeds. (a) Maximum likelihood (ML) phylogenetic tree with Zhongshan partridge duck as the outgroup and 10 migration events. (b) ML phylogenetic tree with Zhongshan partridge duck as the outgroup and 24 migration events. (c) Maximum likelyhood phylogenetic tree with Mallard (Anas platyrhynchos) as the outgroup and 6 migration events. (d) ML phylogenetic tree with Mallard (Anas platyrhynchos) as the outgroup and 30 migration events. ZS, Zhongshan partridge duck; BJ, Beijing duck; LC, Liancheng white duck; WMA, Wenshi partridge duck; WSB, Wenshi white duck; GX, Guangxi small partridge duck; JX, Jingxi large partridge duck; YY, Mallard (Anas platyrhynchos); BZ, Spot-billed duck (Anas poecilorhyncha); FY, Maple leaf duck; YTG, Cherry valley duck; SX, Shaoxing duck; FH, Fenghua duck; GY, Gaoyou duck; SM, Mountain partridge duck; JD, Jinding duck; PT, Putian black duck; LS, Longsheng emerald duck; DL, Donglan duck; RS, Rongshui Xiang duck; WQ, Wenqiao duck; YL, Yulin partridge duck.

References 21

[1]	Chen SF, Zhou YQ, Chen YR, Gu J (2018) Fastp: An ultra-fast all-in-one FASTQ preprocessor. Bioinformatics, 34, i884-i890.
[2]	Chen ZJ, Wan YL, Zhong WJ, Chen SW, Zhou YH, Shi SJ, Jiang L, Ji PC, Yang ZH, Mao ZX, Mou FS (2021) Development and application of soybean in del markers based on whole-genome resequencing datasets. Journal of Plant Genetic Resources, 22, 815-833. (in Chinese with English abstract)
	[陈正杰, 宛永璐, 钟文娟, 陈四维, 周永航, 石盛佳, 蒋理, 戢沛城, 杨泽湖, 毛正轩, 牟方生 (2021) 基于大豆基因组重测序的InDel标记开发及应用. 植物遗传资源学报, 22, 815-833.] DOI
[3]	Danecek P, Auton A, Abecasis G, Albers CA, Banks E, DePristo MA, Handsaker RE, Lunter G, Marth GT, Sherry ST, McVean G, Durbin R, Group GPA (2011) The variant call format and VCFtools. Bioinformatics, 27, 2156-2158. DOI PMID
[4]	Groenen MAM, Archibald AL, Uenishi H, Tuggle CK, Takeuchi Y, Rothschild MF, Rogel-Gaillard C, Park C, Milan D, Megens HJ, Li ST, Larkin DM, Kim H, Frantz LAF, Caccamo M, Ahn H, Aken BL, Anselmo A, Anthon C, Auvil L, Badaoui B, Beattie CW, Bendixen C, Berman D, Blecha F, Blomberg J, Bolund L, Bosse M, Botti S, Zhan BJ, Bystrom M, Capitanu B, Carvalho-Silva D, Chardon P, Chen C, Cheng R, Choi SH, Chow W, Clark RC, Clee C, Crooijmans RPMA, Dawson HD, Dehais P, De Sapio F, Dibbits B, Drou N, Du ZQ, Eversole K, Fadista J, Fairley S, Faraut T, Faulkner GJ, Fowler KE, Fredholm M, Fritz E, Gilbert JGR, Giuffra E, Gorodkin J, Griffin DK, Harrow JL, Hayward A, Howe K, Hu ZL, Humphray SJ, Hunt T, Hornshøj H, Jeon JT, Jern P, Jones M, Jurka J, Kanamori H, Kapetanovic R, Kim J, Kim JH, Kim KW, Kim TH, Larson G, Lee K, Lee KT, Leggett R, Lewin HA, Li YR, Liu WS, Loveland JE, Lu Y, Lunney JK, Ma J, Madsen O, Mann K, Matthews L, McLaren S, Morozumi T, Murtaugh MP, Narayan J, Nguyen DT, Ni PX, Oh SJ, Onteru S, Panitz F, Park EW, Park HS, Pascal G, Paudel Y, Perez-Enciso M, Ramirez-Gonzalez R, Reecy JM, Rodriguez-Zas S, Rohrer GA, Rund L, Sang YM, Schachtschneider K, Schraiber JG, Schwartz J, Scobie L, Scott C, Searle S, Servin B, Southey BR, Sperber G, Stadler P, Sweedler JV, Tafer H, Thomsen B, Wali R, Wang J, Wang J, White S, Xu X, Yerle M, Zhang GJ, Zhang JG, Zhang J, Zhao SH, Rogers J, Churcher C, Schook LB (2012) Analyses of pig genomes provide insight into porcine demography and evolution. Nature, 491, 393-398. DOI
[5]	Li HF, Xu WJ, Zhu WQ, Xue SM, Chen KW (2008) Genetic diversity and evolution of Fujian domestic duck breeds. Biodiversity Science, 16, 607-612. (in Chinese with English abstract) DOI
	[李慧芳, 徐文娟, 朱文奇, 薛素美, 陈宽维 (2008) 福建省家鸭的遗传多样性及其与野鸭的亲缘关系. 生物多样性, 16, 607-612.] DOI
[6]	Lin RY, Li JQ, Yang Y, Yang YH, Chen JM, Zhao FL, Xiao TF (2022) Genome-wide population structure analysis and genetic diversity detection of four Chinese indigenous duck breeds from Fujian Province. Animals, 12, 2302. DOI URL
[7]	Lin Y, Huang M, Li XJ, Zhang XM, Huang YM, Tian YB, Wu ZP (2023) Uncovering genome-wide copy number variations in 8 duck breeds using whole genome resequencing data. Acta Veterinaria et Zootechnica Sinica, 54, 3700-3709. (in Chinese with English abstract) DOI
	[林燕, 黄敏, 李秀金, 张续勐, 黄运茂, 田允波, 伍仲平 (2023) 利用全基因组重测序数据检测8个鸭品种基因组拷贝数变异. 畜牧兽医学报, 54, 3700-3709.] DOI
[8]	Liu Z, Sun H, Lai W, Hu M, Zhang Y, Bai C, Liu J, Ren H, Li F, Yan S (2022) Genome-wide re-sequencing reveals population structure and genetic diversity of Bohai black cattle. Animal Genetics, 53, 133-136. DOI URL
[9]	Luo W (2021) Genetic Diversity of Genome of Local Chicken Breeds in China and Analysis of Selection Imprint of Cockfighting and Antlers’ Crown Populations. PhD dissertation, South China Agricultural University, Guangzhou. (in Chinese with English abstract)
	[罗威 (2021) 中国家鸡地方品种基因组遗传多样性及斗鸡、鹿角冠群体的选择印记分析. 博士毕业论文, 华南农业大学, 广州.]
[10]	Luo W, Luo CL, Wang M, Guo LJ, Chen XL, Li ZH, Zheng M, Folaniyi BS, Luo W, Shu DM, Song LL, Fang MX, Zhang XQ, Qu H, Nie QH (2020) Genome diversity of Chinese indigenous chicken and the selective signatures in Chinese gamecock chicken. Scientific Reports, 10, 14532. DOI PMID
[11]	Mishra S, Sarkar U, Taraphder S, Datta S, Swain D, Saikhom R, Sasmita P, Menalsh L (2017) Multivariate statistical data analysis—Principal component analysis (PCA). International Journal of Livestock Research, 7, 60-78.
[12]	Pickrell JK, Pritchard JK (2012) Inference of population splits and mixtures from genome-wide allele frequency data. PLoS Genetics, 8, e1002967.
[13]	Tubelyte V, Švažas S, Sruoga A, Butkauskas D, Paulauskas A, Baublys V, Viksne J, Grishanov G, Kozulin A (2011) Genetic diversity of tufted ducks (Aythya fuligula, Anatidae) in Eastern Europe. Open Life Sciences, 6, 1044-1053. DOI URL
[14]	Wang R, Sun JL, Han H, Huang YF, Chen T, Yang MM, Wei Q, Wan HF, Liao YY (2021) Whole-genome resequencing reveals genetic characteristics of different duck breeds from the Guangxi region in China. G3: Genes, Genomes, Genetics, 11, jkab054.
[15]	Wang TM, Guo QX, Bai H, Chang GB, Chen GH (2021) Analysis of population structure of different duck genetic resources based on whole-genome resequencing. Chinese Journal of Animal Science, 57(11), 78-81. (in Chinese with English abstract)
	[王统苗, 郭其新, 白皓, 常国斌, 陈国宏 (2021) 基于全基因组重测序对不同鸭遗传资源进行群体结构分析. 中国畜牧杂志, 57(11), 78-81.]
[16]	Weir BS, Cockerham CC (1984) Estimating F-statistics for the analysis of population structure. Evolution, 38, 1358-1370. DOI PMID
[17]	Wu PX, Wang K, Yang Q, Zhou J, Chen DJ, Liu YH, Ma JD, Tang QZ, Jin L, Xiao WH, Lou PE, Jiang AN, Jiang YZ, Zhu L, Li MZ, Li XW, Tang GQ (2019) Whole-genome re-sequencing association study for direct genetic effects and social genetic effects of six growth traits in large white pigs. Scientific Reports, 9, 9667. DOI PMID
[18]	Zhang L (2019) Genome Sequencing and Research of Bama Mini-Pig. PhD dissertation, Guangxi University, Nanning. (in Chinese with English abstract)
	[张笠 (2019) 广西巴马小型猪全基因组测序及分析研究. 博士学位论文, 广西大学, 南宁.]
[19]	Zhang Y (2014) Genetic Diversity and Population Structure Analysis of Some Local Duck Breeds in China. PhD dissertation, Yangzhou University, Yangzhou, Jiangsu. (in Chinese with English abstract)
	[张扬 (2014) 我国部分地方鸭品种遗传多样性与群体结构分析. 博士学位论文, 扬州大学, 江苏扬州.]
[20]	Zhang YH, Jia HX, Wang ZB, Sun P, Cao DM, Hu JJ (2019) Genetic diversity and population structure of Populus yunnanensis. Biodiversity Science, 27, 355-365. (in Chinese with English abstract) DOI URL
	[张亚红, 贾会霞, 王志彬, 孙佩, 曹德美, 胡建军 (2019) 滇杨种群遗传多样性与遗传结构. 生物多样性, 27, 355-365.] DOI
[21]	Zhang ZB (2018) Study on Selection Signal and Domestication Time of Domestic Ducks by Genome-wide Resequencing. PhD dissertation, China Agricultural University, Beijing. (in Chinese with English abstract)
	[张泽宾 (2018) 利用全基因组重测序研究家鸭驯化的选择信号及驯化时间. 博士学位论文, 中国农业大学, 北京.]