The influence of lipid-extraction on the δ13C of mesopelagic and demersal fish in the South China Sea: modification and application of lipid normalization models
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Abstract: Mesopelagic fish, the most important daily vertically migrating community in the oceans, are characterized by high lipid content which may obscure the interpretation of stable isotopes analysis. Demersal fish, which are important consumers in the food web dominated by mesopelagic fish, also have a high lipid content. Here we collected 127 fish samples from the South China Sea and evaluated the effect of lipid contents on δ13C of mesopelagic and demersal fish. In lipid-extracted mesopelagic fish, the C/N content ratio (<5.5) shows a clear correlation with Δδ13C (the offset of bulk and lipid-extracted δ13C values), especially in non-migratory and semi-migratory species; these values were less correlation in demersal fish. Based on our results, we suggest that mesopelagic and demersal fish in different regions of the South China Sea should be studied separately using appropriate correction models and less fit for the traditional model. Moreover, the C/N content ratio should be used cautiously for establishing the lipid normalization model, especially for the fish in migratory mesopelagic fish and demersal fish. Our results also reveal that mesopelagic fish across nearby regions could be analyzed together. The new models described here can be applied in future studies of mesopelagic and demersal fish in the South China Sea.
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Key words:
- mesopelagic fish /
- demersal fish /
- lipid normalization model /
- C/N content ratio /
- lipid content /
- δ13C /
- South China Sea
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Figure 1. Map of sample collection regions in the South China Sea (SCS) (a): the north slope of the SCS (b); the mid-western SCS (c). L1–L10 represent sampling sites of mesopelagic fish on the northern slope of the SCS; D1 and D2 are sampling sites of demersal fish in the continental shelf area and shrimp farm zone of the SCS; S1–S6 represent sampling sites in the mid-western SCS.
Figure 2. The difference between Δδ13Cextraction and Δδ13Ctheory in four groups of fish in the SCS. Δδ13Cextraction: the difference between bulk and lipid-extracted δ13C. Δδ13Ctheory: Δδ13C was calculated using the lipid normalization model (Post et al., 2007). Grey area: ±0.2‰ error range.
Figure 3. The percentage of lipid content, total nitrogen (TN), and total organic carbon (TOC) in mesopelagic and demersal fish in the South China Sea (SCS).
Figure 4. The linear relationship between C/Nmass and Δδ13C of mesopelagic fish in the northern (a), mid-western (b), and demersal fish (c) in two sites of the South China Sea (SCS). Grey area: 95% confidence interval.
Figure 5. The non-linear relationship between C/Nmass and Δδ13C of mesopelagic fish in the northern, mid-western and demersal fish in the South China Sea (SCS).
Table 1. Sample information
Sampling period Region Habitat Site Number Oct. 2014 north slope mesopelagic L1, L4, L5 14 Jun. 2015 north slope mesopelagic L2, L6, L8 32 Mar. 2017 north slope mesopelagic L3, L7, L9, L10 19 Oct. 2016 mid-west mesopelagic S1−S6 24 Oct. 2014 Site D1 demersal D1 24 Jun. 2015 Site D2 demersal D2 14 
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Table 2. Mean±SD lipid content, carbon content to nitrogen content ratios (C/Nmass), total nitrogen content (TN), total organic carbon content (TOC), and Δδ13C (the difference between bulk and lipid-extracted δ13C) in four regions of the SCS. Pearson correlation coefficient tested the correlation between lipid content and C/Nmass, C/Nmass and Δδ13C
Region Lipid content/% C/Nmass TN/% TOC/% Δδ13C/‰ Lipid content vs. C/Nmass C/Nmass vs. Δδ13C P Pearson corr. P Pearson corr. North slope 30.4±6.6 3.7±0.6 10.5±1.0 40.7±5.0* 0.9±0.7* < 0.01 0.38 < 0.01 0.77 Mid-west 31.7±5.9 3.7±0.6 9.2±1.0* 32.9±3.2* 1.2±0.6 0.69 −0.09 < 0.01 0.95 Site D1 16.7±6.9 3.5±0.4 12.8±2.1 43.8±5.9 0.4±0.2 < 0.01 0.86 < 0.01 0.84 Site D2 24.5±6.8 3.1±0.2 12.1±0.3 37.5±1.6 0.9±0.2 0.06 0.53 0.03 0.57 Note: *represents a significant difference in ANOVA for different migration habits in mesopelagic fish with significant value P<0.05.
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