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NOT TO BE CITED WITHOUT PRIOR REFERENCE TO THE AUTHORS ICES CM 2007/D:22 Comparative Marine Ecosystem Structure and Function:

Descriptors and Characteristics Structure and function of three marine ecosystems in Korea: A comparative study by

Chang Ik Zhang Department of Marine Production Management Pukyong National University Busan, 608-737, Korea cizhang@pknu.ac.kr Jae Bong Lee* Fisheries Resources Research Team National Fisheries Research and Development Institute Busan, 609-905, Korea * Corresponding: Alaska Fisheries Science Center 7600 Sand Point Way NE, Seattle, WA 98115 USA jbonglee@gmail.com, leejb@nfrdi.re.kr Tel: (206) 526-4727 Fax: (206) 526-6723 Sun Kil Lee South Sea Fisheries Research Institute National Fisheries Research and Development Institute Yeosu, 556-820, Korea Bernard A. Megrey National Marine Fisheries Service Alaska Fisheries Science Center 7600 Sand Point Way NE Seattle, WA 98115 USA

Abstract
Comparative analyses are a power way to evaluate similarities and differences in ecosystem structure and function. We identified key areas of differences and similarities in the structure and function of three marine ecosystems, that is, East/Japan Sea (ES), Yellow Sea (YS) and East China Sea (ECS) in Korea. Common modeling approaches, such as Ecopath models to three marine ecosystems were applied with important commercial fisheries, and set of macrodescriptor metrics from systems ecology were used to examine large scale ecological characteristics for the three ecosystems. Macrodescriptor metrics were calculated with Ecopath diet matrices, which were provided by food web information in three marine ecosystems. Food web connectance showed that average number of trophic steps from primary producers to apex predators was longest in the EJS, and that trophic pathways were more linear in the YS. ES and ECS had pelagic and benthic foodwebs, while YS mainly pelagic. Other macrodescriptor metrics, such as linked pathways starting from a group and returning to it, were described to compare ecological attributes for the three marine ecosystems.

1. Introduction
There are three offshore zones in Republic of Korea (ROK) waters, consistent with the three large marine ecosystems (LME) around the Korean Peninsula (Fig. 1). Resource management decisions in the offshore zones are made by the central government of the ROK. Management in offshore zones is supported by scientific research by regional institutes of the National Fisheries Research and Development Institute (NFRDI), in the form of regular surveys of oceanographic and biological variables and quantitative assessments of fishery resources. The East Sea (mean depth = 1700 m) is bound by the ROK, the Democratic Peoples Republic of Korea (DPRK), Russia (mainland and Sakhalin Island) and Japan (Fig. 1). ROK waters in the Japan/East Sea are strongly influenced by the North Korea Cold Current, part of 2

the southerly flowing Liman Current that originates in the Sea of Okhotsk; and by the Tsushima warm current, which moves north through the Korea/Tsushima Strait and causes upwelling along the east of the Korean Peninsula. These currents meet near the 40N parallel, creating a strong frontal region (Rebstock and Kang 2003). The Korea/Tsushima Strait connects the Japan/East Sea to the East China Sea. The NFRDI divides these seas along a straight line between Ulgi Lighthouse (southeast coast of ROK) and the southwest tip of Honshu (Japan). The remainder of this offshore zone is delineated by the extent of the EEZ. Additionally, some waters in the central basin are fished under joint agreement

between the ROK and Japan. The East China Sea is bound by the ROK to the north, Japan (Kyushu and the Ryukyu Islands) to the east and southeast, the PRC to the west, and the island of Taiwan to the south; its eastern and southern extents connect to the Pacific Ocean and South China Sea through several wide straits (Fig. 1). To the north-northwest is the Yellow Sea, bound by the ROK, DPRK, and PRC. The shallow Yellow Sea (mean depth = 44 m) lies entirely over the continental shelf, while the East China Sea is deeper (mean depth = 270 m) and somewhat more variable, with 81% of its area over the shelf, 11% over continental slope, and the remainder over a deep basin. The NFRDI line between the East China and Yellow seas extends from the island of Jin-do (near the southwest tip of the Korean Peninsula) to Chagui Island (near Jeju-do) and then across to the mouth of the Yangtze River (PRC). The offshore zones are also delineated by EEZ boundaries. There are also jointly fished waters in the East China Sea (with Japan) and a provisional joint-fishing zone (with the PRC) in the East China and Yellow seas.

Recent arguments for Ecosystem-Based Fisheries Management means moving from single-species population assessments to ecosystem community-based assessments paying particular attention to community dynamics and species interactions (Megrey and Aydin, in review). Comparative analyses are a power way to evaluate similarities and differences in ecosystem structure and function, especially to quantify the impacts of climate variability on trophic transfer and ecosystem structure. We identified key areas of differences and similarities in the structure and function of three marine ecosystems, that is, East Sea (ES), East China Sea (ECS) and Yellow Sea (YS) in Korea. The objectives of this study are to use a common modeling approach, such as an Ecopath model, apply it to the three marine ecosystems with important commercial fisheries, derive a set of system level ecological macrodescriptor metrics to represent meaningful attributes of marine ecosystems, identify key areas of differences and similarities, and to compared our results to published results from other systems.

2. Materials and Methods


2.1 Data sets The three main datasets for this study are taken from published Ecopath models. Zhang et al. (2004) provides an Ecopath model for the East Sea (ES) containing 103 species or functional groups (not including detritus group) covering 4 trophic levels (1.0-4.35) based on data from 1990-1995. Zhang et al. (2006) provide an Ecopath model for the East China Sea (ECS) containing 138 species or functional groups (not including detritus group) and covering 4 trophic levels (1.0-4.18). Zhang and Lee (in review) provide an Ecopath model

for the Yellow Sea (YS) containing 108 species or functional groups (not including detritus group) and covering 4 trophic levels (1.0-4.55). The Ecopath model (Christensen and Pauly, 1992; Pauly et al., 2000), which creates a static mass-balanced snapshot of the resources in an ecosystem and their interactions, represents the ecosystem as trophically linked biomass pools. The biomass pools consist of a single species, or species groups representing ecological guilds. Ecopath data requirements are relatively simple, and generally already available from stock assessment, ecological studies, the literature, biomass estimates, consumption estimates, diet compositions, and fishery catches.

2.2 Ecopath with Ecosim (EwE) Metrics The EwE software provides several system level calculations and metrics that help characterize the ecosystem. A selection of these is briefly described below and more information and details can be found in Christensen et al. (2004).

2.2.1 Summary Statistics (a) Total System Throughput: The sum of all flows in a system expressed as a flow in units of mtkm-2yr-1and represents the size of the entire ecosystem in terms of flow (Ulanowicz, 1986). It is the sum of total consumption, total export, total respiration, and total flow to detritus. (b) Total Primary Production: Total net primary production from all producers expressed as a flow in units of mtkm-2yr-1.

2.2.2 Network Analysis (c) Finns Cycling Index: developed by Finn (1976) it expresses the fraction of an ecosystems throughput that is recycled. This metric is assumed to increase for more mature ecosystems. (d) Finns Mean Path Length: after Finn (1980), the average number of groups that an inflow passes through. This metric is assumed to increase for more mature ecosystems.

2.2.3 Cycles and Pathways (e) Cumulative Number of Pathways from Primary Producers to a Trophic Level: Originally suggested by Ulanowicz (1986), it is the number of pathways implied by the food web representing the ecosystem that connects the primary producers to the higher trophic levels. (f) Sum of flows to detritus: sum of all flows to detritus from all ecosystem components. (g) Total primary production/total respiration: The ratio between total primary production and total respiration in a system is considered to be an important ratio describing the maturity of an ecosystem (Odum, 1971). In the early developmental stages of a system, production is expected to exceed respiration, leading to a ratio greater than 1 however, in mature systems, the ratio should approach 1. (h) Throughput cycled (including detritus): the sum of biomass cycled through the ecosystem, including detritus, in units of mtkm-2yr-1

Results
Figures of the trophic connections from Ecopath (Fig. 2) show some fundamental structural differences in the three ecosystems. Even though both systems cover the same trophic level range, 1-5, the East China Sea appears to have a broader lower trophic base and the East Sea is wider at the upper trophic levels compared with the Yellow Sea. Based on the trophic size-shifted longevity, which is calculated biomass per production of the ecosystem (Fig. 3), the Yellow Sea has a steep slope (0.518) of longevity compared with the East Sea (0.350) and East China Sea (0.402). A summary and highlights of metrics from EwE (Table 1) show the ES and ECS have more throughput, more primary production and less detritus, compared with the YS. The BS ecosystem appears to be more mature than the ES and ECS based on the ratio of total production to total respiration. Finns cycling index shows the YS as the more mature of the three ecosystems, and ES and ECS have longer mean path lengths compared with the YS. The cumulative numbers of paths leading from primary producers to upper trophic levels are almost eight times higher in the ES and YS compared with the ECS. The total amount of primary production flowing to upper trophic levels is over eight times higher in the YS compared with the ES and ECS. We can see a different representation of the relationship between trophic pathway and trophic level in Figure 4. The top panel shows that the cumulative number of pathways from primary producers for the ES and YS increases faster than the ECS at trophic levels 3.5-4.5. The data, plotted on the log scale both show a linear increasing trend, but the slopes are not significantly different from each other (p<0.1).

Discussion
Results from this analysis do not consider any factors that would contribute to uncertainty. Moreover, the food habits data are isolated in time, do not reflect temporal changes in abundance and community composition, and often ecosystems lack historic data on temporal and spatial changes in trophic dynamics. Still the diet matrices are comparable between ecosystems because food habits data collection methods used to characterize food web relationships are very standardized. Ecosystem metrics calculated in this study indicate that the ES is a more productive ecosystem. More of this production makes its way to upper trophic level ecosystems components and less is recycled. In conclusion, ecosystem metrics calculated in this study indicate the average number of trophic steps from primary producers to predators is shorter in the YS. The ES and ECS are mainly pelagic. The greater depth and width of the East Sea and East China Sea and the longer water column results in a greater proportion of the sinking flux being re-mineralised by pelagic microbes, reducing the food supply to the benthos. Consequently, more production in the ES and ECS is unused and goes to detritus (ES: 1700m, ECS: 270m, YS: 44m deep). The YS is more efficient at converting primary production into upper trophic level biomass since there are fewer trophic steps and greater throughput cycled both the pelagic and benthic foodwebs.

Acknowledgements
We want to thank to Mr. Man Woo Lee from KICE and Mr. Sang Cheol Yoon from the NFRDI for their support and the NFRDI Stomach lab for their work in the analysis of an immense amount of stomachs that provide the data for carrying out this research.

References
Christensen, V., and Pauly, D, 1992. Ecopath II - a software for balancing steady-state ecosystem models and calculating network characteristics. Ecological Modelling, 61(3-4): 169-185. Christensen, V., Walters, C. J. and Pauly, D, 2004. Ecopath with Ecosim: A Users Guide. Fisheries Center Research Reports, Volume 12(4), University of British Columbia, Vancouver. 154 p. (available online at www.ecopath.org and www.fisheries.ubc.ca). Finn, J. T, 1976. Measures of ecosystem structure and function derived from analysis of flows. Journal of Theoretical Biology, 56: 758-773. Finn, J. T, 1980. Flow analysis of models of the Hubbard Brook ecosystem. Ecology, 31: 562-571. Megrey, B. A., and K. Y. Aydin. In review. A macrodescriptor perspective of ecological attributes for the Bering and Barents Seas. 27pp. Odum, E. P, 1971. Fundamentals of Ecology. W.B. Saunders Co, Philadelphia. 574 pp. Pauly, D., Christensen, V., and Walters, C, 2000. Ecopath, Ecosim, and Ecospace as tools for evaluating ecosystem impact of fisheries. ICES Journal of Marine Science,57(3): 697-706. Rebstock, G.A. and Y.S. Kang. 2003. A comparison of three marine ecosystems surrounding the Korean peninsula: responses to climate change. Progress in Oceanography, 59:357-379. Ulanowicz, R. E, 1986. Growth and Development: Ecosystem Phenomenology. Springer Verlag (reprinted by iUniverse, 2000), New York. 203pp. Zhang, C. I., and M. W. Lee. In review. Variations in the ecosystem structure in the SE Yellow Sea. 13pp. Zhang, C. I., J. B. Lee, Y. I. Seo, S. C. Yoon, S. Kim. 2004. Variations in the abundance of fisheries resources and ecosystem structure in the Japan/East Sea. Progress in Oceanography, 61: 245-265. Zhang, C. I. et al. 2006. Effects of the 1988/89 climatic regime shift on the structure of the East China Sea ecosystem. 40pp.

Table

Table 1. Summary of ecological metrics calculated by the Ecopath model for the East Sea (ES), East China Sea (ECS), and Yellow Sea (YS) Metric (a) Total system throughput (mtkm yr ) (b) Total net primary production (mtkm-2yr-1) (c) Finns cycling index (% of total throughput) (d) Finns mean path length (e) Cumulative # paths connecting primary producers to upper trophic levels (f) Sum of flows to detritus (mtkm-2yr-1) (g) Total primary production/total respiration (h) Throughput cycled (including detritus) (mtkm-2yr-1) (i) Total flows from primary producers to upper trophic levels (mtkm-2yr-1)
-2 -1

ES 18729 9190.57 0.26 1.98 167299 9265.18 51.60 48.94 219

ECS 17061 8374.81 0.15 1.99 21173 8438.13 50.63 25.96 210

YS 3085 177.37 14.12 1.38 166790 721.85 0.148 435.76 1750

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Figure Legends

Figure 1. Map indicating the geographical proximity of marine regions in Republic of Korea (ROK) waters. Inset represents offshore, coastal, and inshore regions proposed by the NFRDI. Figure 2. Trophic connections for the East Sea (top panel), East China Sea (middle panel), and Yellow Sea (bottom panel) Figure 3. Foodweb longevity (Biomass/Production) for the East Sea (top panel), East China Sea (middle panel), and Yellow Sea (bottom panel) Figure 4. The cumulative number of pathways from primary producers to trophic levels 1-5 (upper panel) and the log of the number of pathways from primary producers to trophic levels 1-5 (lower panel along with linear fits, for the East Sea (ES), East China Sea (ECS), and Yellow Sea (YS).

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Figure 1

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East Sea

East China Sea

Yellow Sea Figure 2.

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East Sea

East China Sea

Yellow Sea Figure 3

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180000 Cummulative Number of Pathways from Pri producers 160000 140000 120000 100000 80000 60000 40000 20000 0 1 2 3 Trophic Level 4 5 East Sea East China Sea Yellow Sea

10000000 log (Number of Pathways from Pri producers)


East Sea

1000000 100000 10000 1000 100 10 1 0

East China Sea Yellow Sea East Sea Linear East China Sea Linear Yellow Sea Linear

Trophic Level

Figure 4

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