Construction and analysis of a coral reef trophic network for Qilianyu Islands, Xisha Islands
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Abstract: Qilianyu Islands coral reefs (QICR), located in the northeastern part of the South China Sea, has been affected by human activities and natural disturbance. To characterize the trophic structure, ecosystem properties and keystone species of this region, a food-web model for the QICR is developed using methods involving a mass-balance approach with Ecopath with Ecosim software. Trophic levels range from 1.00 for detritus and primary producers to 3.80 for chondrichthyes. The mean trophic transfer efficiency for the entire ecosystem is 13.15%, with 55% of total energy flow originating from primary producers. A mixed trophic impact analysis indicates that coral strongly impacts most components of this ecosystem. A comparison of our QICR model with that for other coral reef ecosystems suggests that the QICR ecosystem is immature and/or is degraded.
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Key words:
- South China Sea /
- Qilianyu Islands /
- coral reef /
- Ecopath model /
- food webs /
- ecosystem characteristic
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Figure 4. Qilianyu Islands coral reefs Ecopath model plot of functional group niche overlap. Point colors represent geometric mean of “prey overlap index” and “predator overlap index” (color scale to right); functional groups: 4, 7 small carnivorous and herbivorous fish; 14, 15 coral and zooplankton; 11, 13 other echinoderms and crustaceans.
Figure 5. Qilianyu Islands coral reefs model mixed trophic impact analysis. Positive (blue) and negative (red) values of mixed trophic impact index represent positive and negative effects, respectively. LCF, large carnivorous fish; MCF, medium carnivorous fish; SCF, small carnivorous fish; OF, omnivorous fish; CEF, coral-eating fish; HF, herbivorous fish; CTS, crown of thorns starfish; OE, other echinoderms; OM, other mollusca; SBI, small benthic invertebrates.
Figure 6. Keystone index for Qilianyu Islands coral reefs model functional groups. For each functional group, the keystone index (y-axis) is reported against their relative total impact on the trophic web (x-axis). Overall effects are relative to the maximum effect measured; the x-axis scale is between 0.0 and 1.0. The functional groups are ordered by decreasing keystone index; therefore, the key functional groups are those ranking among the first groups. Circles are proportional to the functional group biomass in the system.
Table 1. Qilianyu Islands coral reefs Ecopath model functional groups
Group name Composition of dominant species Chondrichthyes ray, skate, shark Large carnivorous fish Aprion virescens, Pristipomoides filamentosus, Aphareus rutilans, large grouper, etc. Medium carnivorous fish Labridae, Lethrinidae, Priacanthidae, Mullidae, etc. Small carnivorous fish Holocentridae, Apogonidae, Cephalopholis, Epinephelus merra, etc. Omnivorous fish Pomacentridae, Balistidae, etc. Coral-eating fish Scaridae, Chaetodontidae, etc. Herbivorous fish Pomacanthidae, Acanthuridae, Siganus, etc. Turtles Chelonia mydas, Eretmochelys imbricata, Dermochelys coriacea, etc. Crown-of-thorns starfish Acanthaster planci Giant triton Charonia tritonis Other echinoderms urchin, cucumber, brittle star, starsish Other mollusca bivalve, snail, etc. Crustaceans crab & shrimp Coral Pocillopora damicornis, Pocillopora verrucosa, Acropora humilis, Porites lutea, etc. Zooplankton copepoda, planula, juvenile fish, etc. Small benthic invertebrates polychaeta, etc. Macroalgae coralline algae Turf turf Phytoplankton Bacillariopyta, Pyrrophyta, Chrysophyta, Cyanophyta, etc. Detritus particulate organic carbon & dissolved organic carbon Table 2. Estimates of Qilianyu Islands coral reefs ecosystem food intake for trophic groups
No. Prey (predator) 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 1 chondrichthyes 0.01 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 large carnivorous fish 0.05 0.008 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 medium carnivorous fish 0.14 0.1 0.009 0 0 0 0 0 0 0 0 0 0 0 0 0 4 small carnivorous fish 0.12 0.12 0.13 0.03 0.019 0 0 0 0 0 0 0 0 0 0 0 5 omnivorous fish 0.12 0.05 0.06 0.02 0.01 0 0 0 0 0 0 0 0.005 0 0 0 6 coral-eating fish 0.1 0.05 0.04 0.01 0.06 0 0 0 0 0 0 0 0 0 0 0 7 herbivorous fish 0.1 0.19 0.16 0.03 0.05 0 0 0 0 0 0 0 0.005 0 0 0 8 turtles 0.001 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 9 crown-of-thorns starfish 0 0.001 0.001 0 0.001 0 0 0 0 0.15 0 0 0.001 0 0 0 10 giant triton 1×10−5 0.000 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 11 other echinoderms 0 0.06 0.06 0.05 0.05 0 0 0 0 0.4 0 0 0.005 0 0 0 12 other mollusca 0.05 0.1 0.12 0.1 0.1 0 0 0.1 0 0.3 0 0.01 0.05 0 0 0 13 crustaceans 0.12 0.12 0.14 0.15 0.12 0 0 0.05 0 0 0 0 0.012 0 0 0 14 coral 0 0 0 0 0.05 0.4 0.1 0 0.8 0 0.05 0.06 0 0 0 0 15 zooplankton 0 0 0 0.28 0.1 0 0.05 0.1 0 0.15 0.15 0.25 0.2 0.1 0.01 0.1 16 small benthic invertebrates 0 0.05 0.18 0.2 0.1 0 0 0 0 0 0.05 0 0.133 0 0 0 17 macroalgae 0 0 0 0 0.13 0.3 0.3 0.3 0.07 0 0.2 0.2 0.1 0 0.05 0 18 turf 0 0 0 0 0.11 0.15 0.3 0.4 0.03 0 0.15 0.16 0.135 0 0.09 0 19 phytoplankton 0 0 0 0.03 0 0 0.1 0 0.1 0 0.12 0.22 0.154 0.3 0.6 0.15 20 detritus 0 0 0 0.1 0.1 0.15 0.15 0 0 0 0.28 0.1 0.2 0.4 0.25 0.75 Import 0.189 0.150 05 0.1 0 0 0 0 0.05 0 0 0 0 0 0.2 0 0 Sum 1.000 01 0.999 55 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Table 3. Basic Ecopath model functional group input and estimated output parameters
No. Group name TL B/(t·km−2·a−1) P:B/(a−1) Q:B/(a−1) Un. Q Catch EE P/Q NE OI 1 chondrichthyes 3.80 0.097 0.25 4.72 0.20 − 0.189 0.05 0.07 0.41 2 large carnivorous fish 3.57 0.430 0.79 6.64 0.20 0.200 0.723 0.12 0.15 0.30 3 medium carnivorous fish 3.42 3.781 1.00 8.50 0.20 2.500 0.830 0.12 0.15 0.16 4 small carnivorous fish 3.07 1.700 4.00 13.50 0.20 0.800 0.942 0.30 0.37 0.22 5 omnivorous fish 2.84 1.103 4.50 16.30 0.20 1.500 0.981 0.28 0.35 0.40 6 coral-eating fish 2.45 1.415 2.40 20.00 0.20 0.200 0.878 0.12 0.15 0.30 7 herbivorous fish 2.16 8.156 3.00 28.00 0.40 1.800 0.397 0.11 0.18 0.15 8 turtles 2.32 0.020 0.14 3.50 0.20 0.002 0.878 0.04 0.05 0.29 9 crown-of-thorns starfish 2.90 0.485 1.20 5.00 0.20 − 0.300 0.24 0.30 0.20 10 giant triton 3.34 0.002 1.22 4.08 0.20 0.001 0.950 0.30 0.37 0.07 11 other echinoderms 2.26 3.035 2.20 7.80 0.20 0.500 0.786 0.28 0.35 0.21 12 other mollusca 2.33 8.700 2.50 5.60 0.20 6.000 0.955 0.45 0.56 0.24 13 crustaceans 2.46 4.300 3.20 28.00 0.25 0.500 0.904 0.11 0.15 0.31 14 coral 2.13 19.574 3.00 10.00 0.20 − 0.700 0.30 0.38 0.13 15 zooplankton 2.01 3.510 76.00 242.50 0.30 − 0.416 0.31 0.45 0.01 16 small benthic invertebrates 2.10 3.911 12.00 60.00 0.25 − 0.629 0.20 0.27 0.09 17 macroalgae 1.00 22.000 18.00 − − − 0.375 − − − 18 turf 1.00 30.000 25.00 − − − 0.239 − − − 19 phytoplankton 1.00 8.817 231.00 − − − 0.324 − − − 20 detritus 1.00 315.000 − − − − 0.187 − − 0.24 Note: B: biomass; OI: omnivory index; TL: trophic level; P/B: production/biomass; Q/B: consumption/biomass; Un. Q: unassim. consumption; EE: ecotrophic efficiency; P/Q: production/consumption; NE: net efficiency. Values in bold are estimated by the present model. − reprsents no data. Table 4. Distribution of Qilianyu Islands coral reefs ecosystem flow
Trophic level Consumption
by predatorsExport Flow to
detritusRespiration Throughput V 0.070 0.125 0.406 1.196 1.798 IV 1.650 1.220 6.121 15.040 24.030 III 23.730 5.445 56.840 110.500 196.500 II 194.700 7.207 658.500 713.600 1 574.000 I 1 533.000 2 371.000 2 195.000 0.000 6 099.000 Sum 1 754.000 2 385.000 2 916.000 840.400 7 896.000 Table 5. Transfer efficiency of Qilianyu Islands coral reefs ecosystem trophic levels
Source Trophic level II III IV V Producer 12.37 15.53 11.84 11.1 Detritus 13.64 13.73 12.13 10.43 All flows 12.83 14.84 11.94 10.87 Note: Proportion of total flow originating from detritus: 0.45; transfer efficiencies (mTE, calculated as geometric mean for TL II–IV); from primary producers: 13.15%; from detritus: 13.14%; total: 13.15%. Table 6. Comparison of Qilianyu Islands coral reefs ecosystem characteristics with other coral reefs
Area Qilianyu
IslandsHawaii
IslandCocos
IslandDarwin &Wolf
IslandNanwan
BayUvea
AtollCaribbean Sea Cayos Cochinos Media Luna Group number 20 26 31 32 18 25 22 21 Sum of all consumption/(t·km−2·a−1) 1 810 5 332 22 978 8 880 8 373 292 31 013 27 381 Sum of all exports/(t·km−2·a−1) 2 385 520 20 344 16 200 185 81 700 9 779 Sum of all respiratory flows/(t·km−2·a−1) 840 3 477 12 050 5 278 4 629 86 17 096 16 264 Sum of all flows into detritus/(t·km−2·a−1) 2 916 1 700 6 136 2 151 20 115 346 90 17 881 TST/(t·km−2·a−1) 7 952 11 030 41 184 16 652 49 317 909 220 232 71 305 Sum of all production/(t·km−2·a−1) 3 642 − 978 5 235 21 553 325 10 6510 31 684 TLc 2.62 2.59 3.55 3.02 2.40 3.50 3.64 2.97 GE 0.00 4 0.000 09 2.00×10−6 0.001 0.000 36 0.000 15 1.01×10−9 1.20×10−5 Net p.p./(t·km−2·a−1) 3 182.73 3 895.09 4 583.59 3 408.00 20 199.00 265.30 98 796.00 26 043.00 TPP/TR 3.79 1.12 0.38 0.65 4.40 3.10 5.78 1.60 NSP/(t·km−2·a−1) 2 342.30 417.78 −7 466.37 −1 898.00 15 570.00 179.40 81 700.00 9 779.00 TPP/TB 26.30 5.57 2.32 3.64 9.90 19.95 11.82 10.21 TB:TST/(a−1) 0.015 0.06 0.05 0.06 0.04 0.01 0.04 0.04 Total catch/(t·km−2·a−1) 14.003 0.350 0.010 3.841 7.300 0.039 1.00×10−4 0.311 CI 0.33 − 0.17 0.15 − − 0.30 0.26 SOI 0.21 − 0.40 0.32 − − 0.21 0.20 FCI/% 3.64 6.13 6.50 4.75 3.50 − 1.60 6.95 MPL 2.47 − − − − − 6.65 5.65 A:C/% 27.71 31.50 24.80 29.80 − − 47.00 31.00 mTE/% 13.15 − 12.20 10.70 7.80 − − − Note: GE: Gross efficiency (total catch/net p.p.); net p.p.: calculated total net primary production; NSP: net system production; TST: total system throughput; TLc: mean trophic level of the catch; TPP/TR: total primary production/total respiration; NSP: net system production; TPP/TB: total primary production/total biomass; TB/TST: total biomass/total throughput; CI: connectance index; SOI: system omnivory index; FCI: Finn's cycling index; MPL: Finn's mean path length; A/C: ascendency/capacity; mTE: mean transfer efficiencies. −represent no data. -
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