Spatial differentiation of fish functional groups in the Yangtze River

doi:10.17520/biods.2023095

Abstract

Abstract:

Aim: The composition of fish in large rivers exhibits longitudinal variation along the river influenced by environmental gradients. The River Continuum Concept revolutionized the understanding of river ecosystems by linking changes in river macro-invertebrate trophic functional groups to the differentiation of nutrient sources between upstream and downstream areas. The concept offers a novel framework for studying the distribution patterns of river biomes at large scale, from upstream to downstream. In this study, a dataset encompassing 168 species of fish were used, covering the entire range from Zhimenda, the starting point of the Jinsha River, to the estuary. Functional groups were first classified based on criteria such as body sizes, shapes, feeding habits, and life-history strategies. Subsequently, their distribution patterns as well as their adaptability to environmental factors were investigated across different scales.

Methods: A total of 14 functional groups and 59 combined function groups were classified, and 5 environmental factors were selected: elevation, mean temperature, mean annual temperature range, river width and river slope gradient. The distribution pattern of fish functional groups was analyzed using hierarchical clustering, while ordination analysis was applied to analyze the relationship between environmental factors and fish functional groups at different scales.

Results: The results revealed a primary and secondary differentiation in the distribution of fish functional groups within the Yangtze River: the primary differentiation occurs at Longkaikou, acting as the dividing point, while the secondary differentiation is observed at Shigu, Longkaikou and Downstream of Baihetan Dam. Moving from the upstream to the downstream, fish body sizes transition from small to medium and large, body shapes shift from predominantly fusiform and cylindrical to include compressform appearances. Feeding habits evolve from primarily omnivorous to encompass a more diverse range of feeding functional groups, and life-history strategies transform from opportunistic to periodic and equilibrium strategies. The distribution pattern of fish functional groups is a result of adaptation to spatial differentiation of environmental factors at different scales. Across the larger scale of the entire Yangze River, elevation and temperature, which are associated with climatic features, serve as dominant factors; whereas, as the spatial scale of the study shrinks, the influence of river slope drop associated with topographic features, becomes more prominent and plays the most important role in the I-1 river section at medium scale.

Conclusion: The distribution pattern of fish functional groups is direct outcome of the fish adaptation to environmental differentiation. Furthermore, the specific environment factors that determine the distribution of fish functional groups vary at different scales, consequently affecting the corresponding functional traits of the fish. This study contributes to our understanding of the river continuum theory, the spatial distribution pattern of fish in the Yangtze River, and the environmental adaptation characteristics of fishes.

Key words: the river continuum, the Yangtze River, fish functional groups, spatial differentiation

Anlun Wang, Ping He, Xinyuan Long. Spatial differentiation of fish functional groups in the Yangtze River[J]. Biodiv Sci, 2023, 31(10): 23095.

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URL: https://www.biodiversity-science.net/EN/10.17520/biods.2023095

https://www.biodiversity-science.net/EN/Y2023/V31/I10/23095

Figures/Tables 9

Fig. 1 Landforms of the Yangtze River basin and distribution of fish sampling points. The sites corresponding to the sample points with serial numbers 1?63 are: 1. Zhimenda; 2. Benda Town; 3. Luoxu Town; 4. Maxia Town; 5. Kasongdu Town; 6. Gangtuo; 7. Zengqu River estuary; 8. Jinsha Town; 9. Boluo Town; 10. Yebatan; 11. Zhubalong; 12. Batang River esturary; 13. Suwalong; 14. Benzilan Village; 15. Tongpu Town; 16. Wujing Town; 17. Shangjiang Town; 18. Shigu; 19. Tiger Leaping Gorge; 20. Daju Town; 21. Sanjiangkou Village; 22. Xiashanjiang Village; 23. Heibaishui River estuary; 24. Downstream of Jin’anqiao Dam; 25. Longkaikou; 26. Pianjiao Town; 27. Xintian Village; 28. Jiangbian Village; 29. Yalong River estuary; 30. Yuzuo Town; 31. Jiangtou Village; 32. Upstream of Wudongde Dam; 33. Downstream of Wudongde Dam; 34. Upstream of Baihetan Dam; 35. Downstream of Baihetan Dam; 36. Upstream of Xiluodu Dam; 37. Downstream of Xiluodu Dam; 38. Upstream of Xiangjia Dam; 39. Downstream of Xiangjia Dam; 40. Yibin City; 41. Tuo River estuary; 42. Jiangbei; 43. Jiangjin; 44. Fuling; 45. Wanzhou; 46. Wushan; 47. Gezhou Dam in Yichang City; 48. Qingjiang River estuary in Yichang City; 49. Shishou; 50. Jianli; 51. Jiayu; 52. Wuhan; 53. Huanggang; 54. Hukou County; 55. Wangjiang County; 56. Anqing City; 57. Chizhou City; 58. Tongling City; 59. Wuhu City; 60. Zhenjiang City; 61. Changzhou City; 62. Nantong City; 63. The Yangtze River estuary in Shanghai.

Table 1 Fish functional traits, functional groups and their basis

功能性状 Functional traits	功能群 Functional groups	依据 Basis	参考文献 Reference
体型 Size	小型 Small	最大全长小于20 cm Maximum total length less than 20 cm	Garrison & Link, 2000
	中型 Medium	最大全长在20?50 cm之间 Maximum total length between 20 cm and 50 cm
	大型 Large	最大全长大于50 cm Maximum total length more than 50 cm
形状 Shape	纺锤形 Fusiform	体长 > 体高 > 体宽 Body length > body height > body width	李新辉等, 2022
	圆柱形 Cylindrical	体长 > 体高 ≈ 体宽 Body length > body height ≈ body width
	侧扁形 Compressiform	体长 ≈ 体高 > 体宽 Body length ≈ body height > body width
食性 Feeding habits	碎屑食性 Detritivores	以碎屑、浮游生物和藻类为食 Feeding on plankton, detritus and algae	Iba?ez et al, 2009
	草食性 Herbivores	以水生高等植物为食 Feeding on aquatic higher plants
	无脊椎动物食性 Invertivores	以甲壳类动物、寡毛类动物、软体动物和昆虫为食 Feeding on crustaceans, oligochaetes, mollusks, and insects
	鱼食性 Piscivores	以鱼类为食 Feeding on fish
	杂食性 Omnivorous	以大量植物和动物为食 Feeding on substantial proportions of both plant and animal material
生活史 Life-history	机会策略 Opportunistic	小, 迅速成熟, 寿命较短 Small, rapidly maturing, and short-lived	Winemiller & Rose, 1992; 黎明政, 2013
	周期策略 Periodic	大, 繁殖能力高, 寿命较长 Larger, highly fecund and longer life spans
	均衡策略 Equilibrium	中等大小, 经常表现出亲代照顾, 产生较少但较大的后代 Intermediate size, often exhibit parental care, and produce fewer but larger offspring

Fig. 2 Diagrams of Bray-Curtis similarity clusters of fish functional groups in the Yangtze River. Sample points from upstream to downstream are indicated by numbers 1?63 in order, see Fig. 1 for details. I-1, I-2, II-1, and II-2 are segments of the Yangtze River with reference to the Bray-Curtis similarity of 0.57 and the spatial relationship of the sampling points. Special points refer to the points that are clustered separately of spatially discontinuous with other points of the same cluster.

Fig. 3 Ordination analysis of the distribution of fish combined functional groups in the Yangtze River and the distance from upstream. RDA1 indicates the correlation between them: The combined functional groups with negative values of RDA1 is closer to the upstream; while those with positive values are closer to the downstream. The combined functional groups represented from FG1 to FG59 are shown in Appendix 2.

Fig. 4 Composition of fish functional groups of each secondary segment of the Yangtze River

Table 2 Summary statistic for environmental factors of the Yangtze River at the basin scale and reach scale, and their relationships with the distance from upstream

尺度 Scales	环境因子 Environmental factors	最小值?最大值 Range	平均值±标准差 Mean ± SD	与样点距上游距离的相关性 Relationship with the distance from upstream
尺度 Scales	环境因子 Environmental factors	最小值?最大值 Range	平均值±标准差 Mean ± SD	R	P
流域 Basin	平均气温 Average temperature (℃)	2.65?21.24	15.71 ± 4.62	0.539	**
	平均气温年较差 Average annual range of temperature (℃)	107.00?252.00	180.32 ± 48.68	0.741	**
	海拔 Altitude (m)	0.00?3530.00	1118.51 ± 1140.59	?0.877	**
河段 Reach	河流宽度 River width (m)	60.00?6377.00	829.43 ± 1253.42	0.776	**
河段 Reach	河段坡降 Slope gradient	0.00?0.02	4.8×10^?3± 4.9×10^?3	?0.633	**

Fig. 5 Segmented linear fit equations for the distance from upstream versus average temperature (A), average annual range of temperature (B), altitude (C), river width (D) and slope gradient (E). The segment point of the is the dividing point of the primary divergence of the Yangtze River, Longkaikou (Sampling point 25). All variables were normalized for uniformity of magnitudes.

Fig. 6 Individual importance and total variance explained (R2) of five environmental factors on the distribution of combined functional groups in different spatial scales. A: Whole Yangtze River, segment I and segment I-1 scale; B: Whole Yangtze River, segment II and segment II-2 scale. Segments I-2 and II-1 are not analyzed here due to the small number of sampling points. *** indicates that the total variance explained of the five environmental factors is highly significant (P < 0.001); NS indicates that it is not significant (P > 0.05).

Fig. 7 Spatial patterns of historical and current fish species recording in Yangtze River. * Data from Yang et al (2023).

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