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1 47 2 Vol.47, No OCEANOLOGIA ET LIMNOLOGIA SINICA Mar., 2016 * ( ) 为了解南沙群岛西南部陆架区底层鱼类食物网的营养结构特征, 于 2013 年 9 月利用稳定同位素方法分析了 24 种底层鱼类的碳和氮稳定同位素比值 (δ 13 C 和 δ 15 N 值 ), 并计算了基于 δ 13 C δ 15 N 量化的 6 个营养结构的群落范围指标结果显示 : 南沙群岛西南部陆架区底层鱼类的 δ 13 C 值在之间, 均值为 ( 19.1±0.8) ; δ 15 N 值的范围为 , 均值为 (9.7±1.0) 鱼类的营养位置在之间, 平均值为 3.58±0.29, 表明均以肉食性为主根据群落范围指标结果分析, 鱼类群落的 δ 13 C 和 δ 15 N 值范围 (CR 和 NR) 分别为 3.4 和 4.3, 表明南沙群岛西南陆架区底层鱼类的初始食源较为单一, 营养层次较少 ; 总面积 (TA) 和到重心的平均距离 (CD) 分别为 5.8 和 1.0, 意味着鱼类食物网营养级多样性的总程度和平均程度均较低 ; 平均最近相邻距离 (NND) 及最近相邻距离的标准偏差 (SDNND) 分别为 0.37 和 0.43, 说明研究海域底层鱼类食物网的营养冗余程度较高, 不同种类之间的营养生态位幅宽较小, 重叠程度较高此外, 浅海长尾鲨 (Alopias pelagicus) 和狐形长尾鲨 (Alopias vulpinus) 在南沙群岛西南部陆架区底层鱼类群落的营养结构中起着重要作用因此在对该海域渔业资源进行开发利用时, 应禁止捕捞这些种类, 而对一些冗余种类, 如多齿蛇鲻 (Saurida tumbil) 条尾鲱鲤(Upeneus bensasi) 和带鱼 (Trichiurus haumela) 等, 可以适度捕捞, 以维持底层鱼类群落营养结构的完整性, 在渔业资源合理开发持续利用的同时, 有效地保护海洋生态系统的安全发展南沙群岛西南部陆架区 ; 营养结构 ; 碳和氮稳定同位素 ; 群落范围指标 ; 底层鱼类 Q143 doi: /hyhz ,, (Sheaves, 2006; Heino, 2009),,, (Lindeman, 1942; Bersier et al, 2002; Kaartinen et al, 2012), (Paine, 1980; Polis et al, 1996),,, (Bitterlich et al, 1984; Michener et al, 1994; Gu et al, 1996), *, 2014A ; ( ), 2014TS02 ;, 2013BAD13B06 ;, 2014A ;, NFZX2013,, njpiao1981@163.com :,, feiyanegg@163.com : , :

2 2 : 469 (Fry, 2006),, (Michener et al, 1994) (δ 13 C) (DeNiro et al, 1978; Fry et al, 1984), (δ 15 N) (Cabana et al, 1996; Vander Zanden et al, 1999), Layman (2007a), δ 13 C δ 15 N,,,, (Layman et al, 2007b; Delong et al, 2011; Jackson et al, 2012),, (, 2005) (, 2010), (, 2013),,,,, ( 1), m 92.26m (48.90m), 92.26m, m, 48.90m, 59.40m, 300mm, 50mm, 1h, 3, ( 505μm), 0.7μm (Whatman GF/F) 2h, 60 C 48h,, 1.2 Vario PYRO Cube Isoprime 100, δ : δx( )=(R sample / R standard 1) 1000, X= 13 C 15 N; R= 13 C/ 12 C 15 N/ 14 N; R sample, R standard VPDB( ),, 12 δ 13 C δ 15 N ± (TP) : TP =λ+ (δ 15 N consumer δ 15 N baseline ) / Δδ 15 N, δ 15 N baseline δ 15 N ( λ=1, δ 15 N baseline δ 15 N, λ=2, δ 15 N baseline δ 15 N ), δ 15 N δ 15 N baseline δ 15 N consumer δ 15 N, Δδ 15 N δ 15 N, 3.4 (Post, 2002), δ 13 C δ 15 N, δ 13 C δ 15 N, (Community-wide metrics) (Cornwell et al, 2006; Layman et al, 2007a, 2007b), : (1) δ 13 C (CR): δ 13 C δ 13 C δ 13 C (δ 13 C δ 13 C ) (2) δ 15 N (NR): δ 15 N δ 15 N δ 15 N (δ 15 N δ 15 N ) (3) (TA): δ 13 C δ 15 N (4) (CD): δ 13 C δ 15 N, δ 13 C δ 15 N (5) (NND): δ 13 C δ 15 N

470 47 Fig.1 1 Map of the southwestern continental shelf of Nansha Islands and location of sampling stations (6) (SDNND): δ 13 C δ 15 N Layman (2007a) 2 2.

3 Fig.1 1 Map of the southwestern continental shelf of Nansha Islands and location of sampling stations (6) (SDNND): δ 13 C δ 15 N Layman (2007a) , , 9 9 9, 鲀 2 2 2, 鲉 2 3 3, 1 1 1, 鮟鮟 3 3 3, 1 2 3, cm, (Trichiurus haumela) (Raja kanojei) (Alopias vulpinus) (Alopias pelagicus), 30cm( 1) 2.2, δ 13 C 20.4 ( C. squamiceps) 17.0 ( A. pelagicus), ( 19.1±0.8) ; δ 15 N 7.9 ( C. atridorsalis) 12.2 ( A. pelagicus), (9.7±1.0) ( 1) 2.3 δ 13 C δ 15 N ( 2), : CR NR , TA 5.8, CD 1.0, NND SDNND

4 2 : 471 表 1 南沙群岛西南部陆架区底层鱼类的全长范围 δ 13 C 值 δ 15 N 值和全长的平均值及标准差 Tab.1 Average total length,δ 13 C, and δ 15 N value(average ± SD) of demersal fish species in southwestern continental shelf of Nansha Islands (cm) (cm) δ 13 C ( ) δ 15 N ( ) (n) 1 Champsodon atridorsalis ± ± ± Cheilidonichthys kumu ± ± ± 鲹 Decapterus lajang ± ± ± 鲀 Navodon septentrionalis Lepidotrigla japonica Raja kanojei ± ± ± 鲬 Bembrops caudimacula ± ± ± Citharoides macrolepidotus Upeneus bensasi ± ± ± 鲉 Pterois russelli ± ± ± Malakichthys wakiyai ± ± ± Halieutaea stellata Cubiceps squamiceps ± ± ± Trichiurus haumela ± ± 鲀 Amblyrhynchotus honcheni ± ± ± Synodus macrops ± ± ± Rexea prometheoides ± ± ± Saurida elongata ± ± ± Saurida tumbil ± ± ± 躄 Chaunax fimbriatus ± ± ± 鮟鮟 Lophiomus setigerus ± ± ± Priacanthus macracanthus ± ± ± Alopias vulpinus ± ± ± Alopias pelagicus ± ± ±0.4 2, δ 15 N, 3.58±0.29, (C. atridorsalis), 3.06±0.02; (A. pelagicus), 4.33±0.11 (A. vulpinus) (A. pelagicus) 4.0, ( 3) 3, (, 2003;, 2005;, 2005), 20, (Saulida) (Priacanthus) (, 2005),,,, (, 2003),, 3.1, CR 3.4, (Melville et al, 2005; Vaslet et al, 2012) CR, CR,, (Layman et al, 2007a),,,, CR

5 δ 13 C δ 15 N Fig.2 Stable isotope biplots of δ 13 C and δ 15 N values of the demersal fish species : 1= 2= 3= 鲹 4= 鲀 5= 6= 7= 鲬 8= 9= 10= 鲉 11= 12= 13 = 14= 15= 鲀 16= 17= 18= 19= 20= 躄 21= 鮟鮟 22= 23= 24= Fig.3 3 Mean trophic positions (±SD) of the demersal fish species (Melville et al, 2005),,, (Fanelli et al, 2011; McMeans et al, 2013),, CR NR, NR, (Layman et al, 2007a) NR 4.3, δ 15 N 4.3, δ 15 N 3.4 (Post, 2002), 1.3, (Pauly et al, 2005), 3,, 1.27 Al-Habsi (2008) Nilsen (2008), ,, (Al-Habsi et al, 2008; Nilsen et al, 2008) Pimm(1982),,,,, (, 2014;, 1996), TA CD (Layman et al, 2007a), TA CD , Murdoch (2013) Schmidt (2009), TA CR NR (Layman et al, 2007a), CR NR TA CR NR,, CR ; (Pauly et al, 1998), NR (2003) (2005),,,, (Pauly et al, 1998), NND SDNND

6 2 : 473, NND,,,, SDNND, (Layman et al, 2007a) NND SDNND , (Abrantes et al, 2014; Sabeel et al, 2015),,, (, 2001;, 2005),, ,, (, 2010, 2012),, 3.2 δ 13 C δ 15 N,, δ 13 C δ 15 N,, CR NR TA,, (Layman et al, 2007a), (δ 13 C=( 18.7±0.3), δ 15 N = (10.2±0.3) ) CR NR TA, (δ 13 C=( 18.8±0.2), δ 15 N =(10.2± 0.3) ) 躄 (δ 13 C=( 18.7±0.1), δ 15 N =(10.2± 0.2) ),,, 鲉,,,,,,,,, 4, :, ;,,,,,,, ( ),, 致谢中国水产科学研究院南海水产研究所邱永松研究员在写作过程给予指导, 刘华雪副研究员钟智辉许友伟周艳波杨兵和南锋调查船全体工作人员在样品采集过程提供的帮助和支持, 谨致谢忱,, :, 1 125,,, , 36(6): , , 16(10): 10 19,,, , 37(1): , , 12(5): , , 10(1): 41 45,, , 30(18): ,, , 1(3): 18 25,,, , 12(6): , , 1(4): 18 23,,, , 34(7):

7 474 47,,, 2012., (Trichiurus lepturus)., 43(1): Abrantes K G, Barnett A, Bouillon S, Stable isotope-based community metrics as a tool to identify patterns in food web structure in east African estuaries. Functional Ecology, 28(1): Al-Habsi S H, Sweeting C J, Polunin N V C et al, δ 15 N and δ 13 C elucidation of size structured food webs in a Western Arabian Sea demersal trawl assemblage. Marine Ecology Progress Series, 353: Bersier L F, Banašek-Richter C, Cattin M F, Quantitative descriptors of food web matrices. Ecology, 83(9): Bitterlich G, Gnaiger E, Phytoplanktivorous or omnivorous fish? Digestibility of zooplankton by silver carp, Hypophthalmichthys molitrix (Val.). Aquaculture, 40(3): Cabana G, Rasmussen J B, Comparison of aquatic food chains using nitrogen isotopes. Proceedings of the National Academy of Sciences of the United States of America, 93(20): Cornwell W K, Schwilk D W, Ackerly D D, A trait-based test for habitat filtering: convex hull volume. Ecology, 87(6): Delong M D, Thorp J H, Thoms M C et al, Trophic niche dimensions of fish communities as a function of historical hydrological conditions in a Plains river. River Systems, 19(3): DeNiro M J, Epstein S, Influence of the diet on the distribution of carbon isotopes in animals. Geochimica et Cosmochimica Acta, 42(5): Fanelli E, Cartes J E, Papiol V, Food web structure of deep-sea macrozooplankton and micronekton off the Catalan slope: insight from stable isotopes. Journal of Marine Systems, 87(1): Fry B, Stable Isotope Ecology. New York: Springer, Fry B, Sherr E B, δ 13 C measurements as indicators of carbon flow in marine and freshwater ecosystems. Contributions in Marine Sciences, 27: Gu B, Schell D M, Huang X H et al, Stable isotope evidence for dietary overlap between two planktivorous fishes in aquaculture ponds. Canadian Journal of Fisheries and Aquatic Sciences, 53(12): Heino J, Biodiversity of aquatic insects: spatial gradients and environmental correlates of assemblage-level measures at large scales. Freshwater Reviews, 2(1): 1 29 Jackson M C, Donohue I, Jackson A L et al, Population-level metrics of trophic structure based on stable isotopes and their application to invasion ecology. PLoS One, 7(2): e31757 Kaartinen R, Roslin T, High temporal consistency in quantitative food web structure in the face of extreme species turnover. Oikos, 121(11): Layman C A, Arrington D A, Montaña C G et al, 2007a. Can stable isotope ratios provide for community-wide measures of trophic structure? Ecology, 88(1): Layman C A, Quattrochi J P, Peyer C M et al, 2007b. Niche width collapse in a resilient top predator following ecosystem fragmentation. Ecology Letters, 10(10): Lindeman R L, The trophic-dynamic aspect of ecology. Ecology, 23(4): McMeans B C, Rooney N, Arts M T et al, Food web structure of a coastal Arctic marine ecosystem and implications for stability. Marine Ecology Progress Series, 482: Melville A J, Connolly R M, Food webs supporting fish over subtropical mudflats are based on transported organic matter not in situ microalgae. Marine Biology, 148(2): Michener R H, Schell D M, Stable isotope ratios as tracers in marine aquatic food webs. In: Lajtha K, Michener R H eds. Stable Isotopes in Ecology and Environmental Sciences. Oxford: Blackwell, Murdoch A, Klein G, Doidge D W et al, Assessing the food web impacts of an anadromous Arctic charr introduction to a sub-arctic watershed using stable isotopes. Fisheries Management and Ecology, 20(4): Nilsen M, Pedersen T, Nilssen E M et al, Trophic studies in a high-latitude fjord ecosystem-a comparison of stable isotope analyses (δ 13 C and δ 15 N) and trophic-level estimates from a mass-balance model. Canadian Journal of Fisheries and Aquatic Sciences, 65(12): Paine R T, Food webs: linkage, interaction strength and community infrastructure. Journal of Animal Ecology, 49(3): Pauly D, Christensen V, Dalsgaard J et al, Fishing down marine food webs. Science, 279(5352): Pauly D, Palomares M L, Fishing down marine food web: it is far more pervasive than we thought. Bulletin of Marine Science, 76(2): Pimm S L, Food Webs. Netherlands: Springer, 1 11 Polis G A, Winemiller K O, Food Webs: Integration of Patterns & Dynamics. US: Springer, Post D M, Using stable isotopes to estimate trophic position: models, methods, and assumptions. Ecology, 83(3): Sabeel R A O, Ingels J, Pape E et al, Macrofauna along the Sudanese Red Sea coast: potential effect of mangrove clearance on community and trophic structure. Marine Ecology. 36(3): Schmidt S N, Vander Zanden M J, Kitchell J F, Long-term food web change in Lake Superior. Canadian Journal of Fisheries and Aquatic Sciences, 66(12): Sheaves M, Scale-dependent variation in composition of fish fauna among sandy tropical estuarine embayments. Marine Ecology Progress Series, 310: Vander Zanden M J, Rasmussen J B, Primary consumer

8 2 : 475 δ 13 C and δ 15 N and the trophic position of aquatic consumers. Ecology, 80(4): Vaslet A, Phillips D L, France C et al, The relative importance of mangroves and seagrass beds as feeding areas for resident and transient fishes among different mangrove habitats in Florida and Belize: evidence from dietary and stable-isotope analyses. Journal of Experimental Marine Biology and Ecology, : THE TROPHIC STRUCTURE OF DEMERSAL FISH SPECIES IN SOUTHWESTERN CONTINENTAL SHELF OF NANSHA ISLANDS, SOUTH CHINA SEA NING Jia-Jia, DU Fei-Yan, WANG Xue-Hui, WANG Liang-Gen, GU Yang-Guang, LI Ya-Fang (South China Sea Fisheries Research Institute, Chinese Academy of Fishery Sciences; Guangdong Provincial Key Laboratory of Fishery Ecology and Environment; Scientific Observing and Experimental Station of South China Sea Fishery Resources and Environments, Ministry of Agriculture; Key Laboratory of South China Sea Fishery Resources Development and Utilization, Ministry of Agriculture; Guangzhou , China) Abstract We investigated carbon and nitrogen stable isotope ratios (δ 13 C and δ 15 N) of demersal fish species collected in September 2013 in southwestern continental shelf of Nansha Islands, South China Sea by trawling (the net was 92m long, perimeter 128m, floating line 49m, lead line 59m, mesh size 50mm, trawling time 1h at 3kn/h) to study the trophic structure of the fish species. Several quantitative metrics based on the δ 13 C and δ 15 N ratios were calculated to quantitatively characterize the trophic structure in community-wide aspect. The results show that, the average δ 13 C and δ 15 N values of the demersal fish species in the study region were ( 19.1±0.8) and (9.7±1.0), in range of and , respectively. The average trophic position was 3.58±0.29 and ranged in , indicating that these demersal fish species were mostly carnivorous. δ 13 C range and δ 15 N range values based on the quantitative metrics were 3.4 and 4.3 respectively, suggesting a few basal resources in low trophic level of the fish food web in the study area. Total area and mean distance to centroid values were 5.8 and 1.0 respectively, indicating that the overall and average degree of trophic diversity were low. Mean nearest neighbor distance and standard deviation of nearest neighbor distance values were 0.37 and 0.43 respectively, suggesting that the fish food web in this study area has high trophic redundancy, small trophic niche width as well as low degree of trophic niche separation. In addition, fish species Alopias vulpinus and Alopias pelagicus played an important role in trophic structure of fish community of this area. To maintain the integrity of the trophic structure, ensure the sustainable use of fishery resources, and protect marine ecosystem, we recommend forbidding the capture of these two species and moderately fishing some redundant species (Saurida tumbil, Upeneus bensasi, Trichiurus haumela, etc.). Key words southwestern continental shelf of Nansha Islands; trophic structure; carbon and nitrogen stable isotopes; community-wide metrics; demersal fish

2 : 111 ( Pinctada maxima), 13, 11,,,,,, 15, 25 cm, cm, 4 5 kg 1-2],,, 3-7] 24] 25-26],,, ] min, 1, 5 m 4 m, 20 m , 9-22],,, (

2 : 111 ( Pinctada maxima), 13, 11,,,,,, 15, 25 cm, cm, 4 5 kg 1-2],,, 3-7] 24] 25-26],,, ] min, 1, 5 m 4 m, 20 m , 9-22],,, ( 12 2 2 0 1 6 4 South China Fisheries Science Vol. 12, No. 2 Apr., 2016 doi: 10. 3969/ j. issn. 2095-0780. 2016. 02. 016,,,, (,,, 510300) : ( Pinctada maxima),,,, 54. 55%, 0 0. 25 m - 2, 0 70. 31 gm - 2,